Architecture Follows Nature Biomimetic Principles for Innovative Design - (2023)



Skin is a complex organ that performs a multitude of functions; namely, it serves
as a link between the body and the environment. Similarly, building envelopes
act as interfaces between their inhabitants and external elements. The resulting
architectural designs illustrate an integrative methodology that allows architecture
to follow nature.

“Ilaria Mazzoleni, in collaboration with biologist Shauna Price, has developed a profound
methodology for architectural and design incentives that anticipates and proposes novel
ways to explore undiscovered biological inspirations for various audiences.”

-Yoseph Bar-Cohen

Architecture Follows Nature

Biology influences design projects in many ways; the related discipline is known as
biomimetics or biomimicry. Using the animal kingdom as a source of inspiration,
Ilaria Mazzoleni seeks to instill a shift in thinking about the application of biological
principles to design and architecture. She focuses on the analysis of how organisms
have adapted to different environments and translates the learned principles into
the built environment. To illustrate the methodology, Mazzoleni draws inspiration
from the diversity of animal coverings, referred to broadly as skin, and applies them
to the design of building envelopes through a series of twelve case studies.


Applying Properties of Animals Skins to Inspire Architectural Envelopes

Ilaria Mazzoleni

in collaboration with Shauna Price

Architecture Follows Nature

ISBN: 978-1-4665-0607-7


9 781466 506077

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Architecture Follows Nature
Biomimetic PriNciPles For iNNovAtive DesigN

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CRC Press Series in Biomimetics
Series Editor: Yoseph Bar-Cohen
Jet Propulsion Laboratory, California
Institute of Technology

Architecture Follows Nature—Biomimetic Principles
for Innovative Design
Ilaria Mazzoleni
Biomimetics: Nature-Based Innovation
Yoseph Bar-Cohen
Biomimetics and Ocean Organisms:
An Engineering Design Perspective
Iain A. Anderson and Julian Vincent
Mechanical Circulatory Support for Heart Failure
Pramod Bonde and Robert L. Kormos

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Architecture Follows Nature
Biomimetic PriNciPles For iNNovAtive DesigN

ilaria mazzoleni

in collaboration with shauna Price

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Cover design by im studio mi/la, Richard Molina.

CRC Press
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Version Date: 20130329
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To my parents
& to our green valley

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Foreword by Jannette Yen




Project Credits





Theoretical Framework




  How Nature Inspires Architecture


Historical Interplay of Bio-Inspired
Architecture among Science, Art and Design


   Contemporary Challenges and Interests


   Nature and the Built Environment


  Novel Practices in the Built Environment:
   Dynamic, Atmospheric, and Active


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  How Biology Informs Architecture


  Evolving and Adapting to Survive


  Climate and Biomes


  Ecosystems and Biodiversity


 Lessons Learned: Biology to the
Built Environment






   Skin Composition and Functions


   The Four Selected Functions




  Urania moth (Chrysiridia rhipheus)66
 Violet-tailed sylph (Aglaiocerus coelestis)


  Lettuce sea slug (Elysia crispata)86

Thermal Regulation


 Side-blotched lizard (Uta stansburiana)


 Snow leopard (Panthera uncia)


 Polar bear (Ursus maritimus)




Water Balance

 Banana slug (Ariolimax columbianus)


  Dyeing dart frog (Dendrobates tinctorius)


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  Ochre sea star (Pisaster ochraceus)


 Namib Desert beetles (Onymacris unguicularis,
Physasterna cribripes)




 Tree pangolin (Manis tricuspis)


 Hippopotamus (Hippopotamus amphibius)


Author Biographies



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All around us, we see the ingenious adaptive capabilities in the
form and function of natural systems and the built environment.
From single cells to multicellular organisms to ecological communities, we see architectural parallels in the function and form
of the bivouac tent to the Louvre pyramid to Dubai’s cityscape. We
use the words exquisite and elegant without reserve to describe the
wonders of nature, but can we always apply these superlatives to
our tenements and washed out roads caused by how humans seem
to intrude instead of blend with nature? One solution that raises
our consciousness of the role man plays in the natural ecosystem
of planet Earth is biomimicry. Copying can often be regarded as
high praise and searching for life’s analogies gives hope that perhaps humans may find where they fit by emulating natural interactions and balances, integrating complex functions seamlessly
beneath a semi-permeable skin. The possibilities are endless but
the way ahead is not spelled out. Biologists and non-biologists
alike are becoming entranced by the potential of biologically
inspired design. Yet those who have attempted to teach it note that
there are few guidelines that make this process transparent. For
our next steps, it has become clear that a deep knowledge of biology may serve us better than a superficial skimming that reduces
the complex elegance to mistaken functions. Scientific knowledge
is not that inaccessible anymore, given the new collections such
as Planet Earth that show us some of the most dazzling feats of
nature. Many of the mechanisms of these charismatic organisms
are known and it is with great fun and fascination that today’s

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practitioners of biologically inspired design are diving gleefully
into a treasure chest full of natural solutions waiting to be realized in human design.
To tempt us even further is this treatise by Ilaria Mazzoleni. Be
amazed by the known biomimetic designs that exist, the known
natural systems left to be translated, the exponential rate of technological advances that will enable you, too, to be a bio inspired
designer! The information is at your fingertips but the translation will be made easier by making a friend with a biologist. Go
ahead. Let’s match the complexity of nature with a complexity
of expertise in an interdisciplinary team needed for bio inspired
design. We have the knowledge; we just need more practice using
it. Take a closer look at the life around you and one day you’ll see
much more of it mirrored in the tools we use, the houses we live
in, the rules we rely on. This new intuition gained by using this
approach may actually help us find harmony with nature.

Jeannette Yen
Professor, School of Biology
Director, Center for Biologically Inspired Design
Georgia Institute of Technology, Atlanta, Georgia

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I am deeply connected to two places in this world: Los Angeles,
California — with its expansive horizon, deep ocean, blue sky,
desert light, and sunny days; and Val Taleggio, Italy – with its
immersive green landscape, deep mountainous canyons, white
peaks, and fresh air. These natural environments have shaped my
profound love for nature, but most importantly have connected
me to the influential people who have molded me into the person
I am today. My deepest appreciation goes to them for supporting the genesis and development of this book and celebrating its
completion. Without them nothing would have been meaningful
to begin with, and because of them I hope to modestly contribute
in shaping a better world.
First and foremost, I am truly grateful to Shauna Price; without
her contribution the book would not have been possible. Our
collaboration started with her participation in my biomimetic
courses, where she instilled fundamental concepts which have
been refined further in the writing of this book. She kindly nurtured my interest in learning about the governing principles of
nature and welcomed the idea of a collaboration between biology
and architecture. Fantastic and nourishing ideas have flourished
with help from our team at im studio mi/la: Richard Molina’s
dedication has been instrumental with the management and
graphic integrity of the book’s complex and rich material. A special thank you to Juan Miguel San Pedro for his positive energy in
editing all diagrams and drawings of the twelve projects included
in the book.

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I would like to express my gratitude to Janine Benyus and Steven
Vogel for their positive feedback on preliminary drafts of the book
which encouraged me to show it to Yoseph Bar-Coehn. Yoseph’s
immediate enthusiasm in wanting to collaborate as the editor and
proposing the book to CRC Press has been a very gratifying experience. I am thankful to Jeannette Yen for her kind words in the
Since 2005, the Southern California Institute of Architecture
(SCI-ARC) and the institute’s directors, chairs and coordinators
have all graciously supported the notions and implementation of a
variety of biomimetic seminars I have proposed and taught. Over
these years various colleagues and interdisciplinary collaborators have participated in the seminars through a series of lectures and critiques which have created a nurturing, collaborative
atmosphere which we all benefited and grew from. Thank you
to the following people: Berenika Boberska, Juan C. Portuese,
Yo Yoshima, William Howard, Lorraine Lin, Steve Ratchye,
Prof. Geoff Spedding, Borja Mila, Jaime Chavez, and a number
of other biologists who specifically contributed to the editing of
Part II.
Different segments of the manuscript have found the keen eyes
of readers and editors such as architect Sarah Graham and environmental lawyer Suzie Lieberman. The patient guidance and
beneficial input provided by Sarah Dennison has been instrumental through the process; being both a biologist and architect, no
one better could interpret my thoughts and guide me through the
maze (and my obfuscation) of the English language.
Useful ideas and clarifications emerged from conversations with
Alessandro Mendini, Stefano Boeri, Pam Thompson, Mohamed
Sharif, Walid Soussou, Deborah Gloria, Diego Terna, Asli Suner,
Jan Ipach, Claudio De Fraja, Massimo Garbuio, Fabrizio Gallanti,
Jeffrey Landreth, Anne McKnight, and Arun Krishnan. Throughout
the process they graciously supported me, encouraged me, and aided
with brainstorming sessions.
The vibrancy of the book lies not only in its text, but also in
the carefully selected imagery. The team early on decided to
find images of animals photographed by scientists. The long
and intense search for these images became a journey all on
its own, allowing us to become acquainted through email with

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great people dedicating their life to field work. It has been an
adventure to reach out, learn what they are doing, and encounter
such generosity in sharing their photographs. I truly loved this
process – virtually traveling with them, immersing myself in their
observations, and validating that great discoveries are still ongoing in the twenty-first century. Thank you all!
I would like to conclude by thanking my family and friends, all of
whom have supported my desire to write this book for years and
continuously encouraged me forward.
I wish the reader a pleasant time of discovery!

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Project Credits 

I gratefully offer thanks to the students who enthusiastically contributed to the book, in particular those who put extra effort into
their original seminar project. You know who you are!
Most of the projects were undertaken by SCI-Arc students and
initiated under various designations of the seminar “Biomimicry:
Innovation in Architecture Inspired by Nature” throughout the
academic years of 2010–2011. An important and fundamental role
is attributed to the interdisciplinary editorial team, who have read,
analyzed and edited the projects – transcending their strengths
and coherency. To each and all of them, my profound appreciation, in particular biologists Graham Slater, Ryan Ellington, Ryan
Harrigan, and architect Stacy Nakano. Thank you.

Project Team Members
Urania moth – Benedetta Frati, Nir Zarfaty
Violet-tailed sylph – Joanna-Maria Helinurm, Alina Amiri
Lettuce sea slug – Ana Munoz, Ryan Hopkins
Side-blotched lizard – Juan Miguel San Pedro, Alex
Nahmgoong, Yuan Yuan
Snow leopard – Joakim Hoen, Mamoune Ghaiti
Polar bear – im studio mi/la, Ilaria Mazzoleni, Alessandro
Colli, Richard Molina
Banana slug – Astri A. Bang, Maya Alam, Janni S. Pedersen
Dyeing dart frog – Erin Lani, Jordan Su
Ochre sea star – Paul Mecomber, Adrian Ariosa
Namib desert beetles – Emily Chen, Carlos Rodriguez
Tree pangolin – Ross Ferrari, Thomas Carpentier
Hippopotamus – Sarah Månsson, Worrawalan Raksaphon

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Biology is a rich discipline. Its influence spreads far beyond the
field of science and extends into seemingly unrelated fields like
architecture. The term “biomimetics” has been in use since the
1960s when Otto H. Schmitt defined it as “biology + technology”
but applied it mainly within the field of engineering. In the field
of architecture, however, biomimicry has only been used since
the early 2000s, reconsidering biomimetics as applied to design.1
While biomimicry has been typically limited to imitating the morphological aspects of the biological world, its potential to reveal
the functional aspects has been largely overlooked. This book, by
providing an understanding of fundamental biological concepts
of evolution and, in particular, adaptation, proposes a new way of
looking at the functional aspects of nature as sources of inspiration
for improvements in application to building design and form.
In this book the biomimetic approach to design is seen as an opportunity to bring attention to the unique capacity of nature to work
in a systemic way. In fact, everything in nature is interconnected
to the degree that some scientists consider Earth a single ecosystem or biosphere discouraging discrete views of smaller scale systems that exhibit discontinuity and separation. Nature’s systems
are dynamic, in flux, and in constant transformation, subject to
the laws of physics. Interconnectivity also inherently involves the
concepts of coexistence and richness in biodiversity. Natural systems depend upon the diversity of their elements, which ultimately
create balance within an organism or ecosystem. In architecture
we are accustomed to thinking of elements in isolation. And while

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this allows concentration on details, it misses the possibility of
working within a model of ecological, dynamic contextualization.
The study of interconnectivity of individual elements in nature
can lead to a significant shift in contemporary architectural thinking. If inspiration is taken from biology it is easier to understand
nature’s importance and to find meaningful design examples. In
design it is always useful to begin with a simple, holistic building concept. This book is concerned with building envelopes and
their multifunctional connective potential. Building envelopes
may be thought about in terms of both the interior realms of the
building and their capacity to address the forces of the larger
exterior environment. From part to whole, from local to global,
from micro to macro, everything is connected and connectable,
in constant, harmonious flux and unfixed in time or space. From
buildings to communities we can design complex ecosystems rich
in interconnected solutions which offer an infinite range of possibilities. Ecosystems can generate concepts for allowing all elements to coexist with each other. Learning from nature’s efficient
interconnectedness allows designers to consider opportunities for
multifunctional uses and streamlined design solutions. This can
ultimately reduce and rebalance our footprint on Earth while producing an integrated built and natural ecosystem, eliminating the
separation between artificial and natural. Moving from designing
buildings in isolation to considering them within larger networks
involves a simultaneous understating of a multitude of factors.
Part of this transformative shift in thinking also involves reimagining the role of architecture in terms of its restorative potential.
Restoration of natural systems could become a significant contribution of architecture.
The case studies presented in the book are explorations of the animal kingdom that consider the climatic and ecological contexts
in which the selected animals evolved. Adaptations, which allow
animals to survive in their habitats, provide lessons that, in turn,
are translated into designs for the built environment. Such investigations focus on the analysis of various animal skins and skin
coverings. Animal skins are one of the major systems for which
architecture can draw inspiration from biology. There are, however, many others, such as plant species, that might be considered,
and the methodology proposed in this book can be used to guide
one through the process in an analogous fashion. Skin is a complex and remarkably sophisticated and variable organ, providing
animals with protection, sensation, and heat and water regulation.

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The term “skin” is used in this book on a general level to refer to
any animal covering, including fur, feathers, scales, exoskeletons,
and shells. Skin is understood as an interface, transcending its
surface, giving the appearance of something that separates, but
instead acts as a threshold or boundary, allowing for interaction
with the elements in multiple directions, scales, and timeframes.
Similarly, building enclosures provide building inhabitants an
interface with environmental elements such as weather, noise,
and sunlight. Architectural enclosures furnish great opportunities to take into consideration dynamic local environmental conditions, creating the potential to use the conditions as resources
to be enhanced and supported rather than simply as elements to
conceal or overcome. Biomimicry is a rich tool for designers to
innovatively integrate the local environment into their projects,
supporting a more sustainable way of building and living.
In its use to date, biomimicry has been predominantly applied
to form in architecture. Instead, this book is interested in focusing on biomimetic applications to architecture in both form
and function. The methodology is deployed through a series of
sample projects that resulted from a collective investigation that
took place during the course of a seminar born from ongoing professional research by the author and her academic collaboration
with students at the Southern California Institute of Architecture
and biologists from the Department of Ecology and Evolutionary
Biology, University of California, Los Angeles.
The book is organized into two parts. Part I introduces and
describes the principles of design that take inspiration from
nature as well as the fundamental biological concepts that can
inform architecture. The biomimetic methodology, developed by
the author, is introduced in Part II which explores four sets of case
studies, each of which investigates a particular function of skin:
communication, thermoregulation, water balance, and protection. The integration of these functions with other internal body
systems is also explained. The chapter on each animal has two
components. The first introduces and analyzes a selected animal
and its skin functions. The second assimilates biomimetic applications within a theoretical “proto-architectural” project, where
the prefix proto- indicates that the projects here presented are at
their conceptual stage of development.2 Proto-architectural thus
defines a key moment of design, open to discourse, experimentation, and environmental innovation. The examples are intended as

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a springboard for applying the functional analysis of an animal’s
biology to the design process.
The new methodology provides a path for drawing design inspiration from nature. It considers architecture beyond the aesthetic
or functional, and begins to explore the conceptually strategic.
The book proposes a novel way of looking at the environment and
provides cues to undiscovered inspirations for a variety of audiences: for designers and architects, it provides biological inspiration for further flexible and dynamic design, and templates for
them to derive other species examples, and acts as a springboard
for exploring broader applications beyond envelopes to applying whole ecosystems to entire buildings and communities. For
engineers, it is hoped that the book might inspire new building
technologies and design of materials. The book can be used by
biologists to continue researching nature and to make their findings available and understandable to designers and architects. For
environmentalists, it may be used to help overturn the stereotype
that ongoing building construction must inherently be ecologically damaging, neither capable of maintaining nor restoring land.
For others, the book looks to promote greater appreciation of biodiversity and incentives for conservation by bringing attention to
the vast variety of species and their adaptation to their environment. Architecture Follows Nature—Biomimetic Principles for
Innovative Design seeks to inspire a shift in thinking about the
built and unbuilt worlds by illustrating that they can be intelligently integrated, and that bio-inspired design might not only help
the environment but also be beautiful!

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  Theoretical Framework


“Nature does nothing uselessly.”

- Aristotle

What is nature? What is our relationship with nature? How can
we change our relationship with nature to mitigate the impacts we
have brought upon its systems and ourselves?
The word “nature” has its roots in the Latin word natura, which
in the classical era meant “birth” or “begetting.” “Nature” is also
conceptually close to the Greek word physis, which refers to a
material system that exists and grows. Accordingly, nature is
by no means static and is constantly regenerating and transforming itself. The interconnectedness of natural systems on Earth
supports life, which evolves at myriad speeds and scales so far
not found anywhere else in the universe. Interconnectedness is
the theoretical concept for this book, defined as the integration
of nature’s solutions with innovative problem solving for manmade environments. The Earth’s natural resources are finite and
the planet is increasingly vulnerable to human activities, making
apparent the responsibility we have to lessen our impact on it.
The process of evolution and the resulting adaptations have
allowed life to sustain itself for millenia. But the increased pace
and scale of human activities has unknown consequences for the

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balance of systems that allow all species, including our own, to
thrive. Sustainable design is a way for us to begin to harmonize
man-made structures with the natural environment. Biomimicry
can help us change our perception by looking to nature as a source
of functional and aesthetic solutions rather than as a source of
obstacles to overcome.
Technological developments have generally been born from
the mindset that nature can be harnessed to suit human needs.
Historically, humans have used ingenuity to develop tools that
have allowed us to transcend the environmental stresses threatening our survival. From the spear to the wheel to the building, our
inventions have allowed us to catch prey, travel further distances,
and create shelter in otherwise inhospitable places. Architectural
design has grown out of the need for shelter and expression, and
today we have the ability to design and engineer buildings with
technologies and strategies that provide high levels of comfort
in any climate. Although technology has been very successful in
facilitating our survival, technology has also resulted in the unintended consequences of resource depletion, pollution, and climate
change. During the past few decades it has become clear that a
new way of thinking is inevitable to address these issues.
The application of the life sciences in building design has so far
been limited largely to the imitation of organic form. Biologists
and ecologists have studied the environmental influences on
animal and plant physiology and behavior, but translating these
observations and analyses into architecture has been largely
unexplored. The resilience of species in a particular habitat can
provide valuable lessons for long-lasting design. Just as animals
have systems, such as skeletal, circulatory, immune, digestive,
communication, and sensory, so too do buildings have systems of
structure, circulation, protection, energy and water use, communication, and thermal regulation. Viewed as a network of internal
systems interacting with its surrounding environment, which is in
turn part of a larger global network of systems, the building can
find inspiration from an animal’s interactions with its ecological
This book shows how explorations of the animal kingdom can
help us consider the climatic and ecological contexts in which
the animals selected as case studies have evolved. Building envelopes share much in common with animal skins and can borrow

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an enormous amount of information from them. Like an animal
skin, a building enclosure acts as an interface, allowing for interaction with the elements. Enclosures have the potential to act like
natural filters with the environment, rather than barriers, by being
reactive to the direction of local winds, solar orientation, and
humidity. One can imagine a building that reacts to changes in
weather by altering its shading configuration or water-capturing
abilities. Architectural enclosures, once reimagined as flexible
and reactive, can furnish opportunities for taking into consideration dynamic local environmental conditions that might allow
buildings to co-exist with nature, preventing nature’s degradation
and perhaps contributing to its restoration.
How can architects and designers move beyond the formal imitation of nature to more sophisticated, nature-inspired, performancebased building design? Successful nature-inspired design would
need to include teams of collaborators from multiple disciplines,
not only engineers, contractors and owners/users. Physicists, biologists, ecologists, and other scientists would also be included as
part of the team for their expertise in understanding natural solutions. Biomimicry stresses the interconnectedness of systems to
solve complex problems; similarly, the integration of varied disciplines yields fertile ground for comprehensive designs to address
the array of environmental issues in which our buildings are constructed and operated. Smart solutions derived from examining
nature have the potential to harmonize with the environment,
rather than exploit it.
Architects, as artists, constantly look to new and emerging ways
of exploring and developing ideas to create buildings that not only
function well, but also express the culture and technologies of their
time and set the standard for future ways of living. Contemporary
Parametric Design, for instance, is an area of study where designers manipulate the algorithmically coded parameters of digital
models to organically generate complex geometries, which are
then used for building facades and structures. Furthermore,
nanotechnology has provided the potential to integrate carbon
nanotubes in building materials to create surfaces that react to
our touch. Through experimentation, imagination and creativity,
biomimetic design is a new direction in which to discover and
transform the way we, and our built world, relate to the natural
world. There is a growing movement to RE- : REthink, REduce,
REpair, REuse, REcycle and REimagine the ways in which we

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inhabit our planet. It is the motivation for the studies described
in this book.

How Nature Inspires Architecture
Architecture has always inserted itself into and interacted with
the natural environment. Essentially, architecture provides shelter
in nature to protect its inhabitants from nature.

Human shelters carved out of the
rock in Cappadocia, Turkey, 4th
Century. (Photograph courtesy of A.

Historically, form has been the primary source of inspiration from
nature, ranging from simple formal influences to the more symbolic translation into architectonic language. The Ancient Greeks
fashioned the ornamentation of their columns and temples on
local plant life to symbolize nature. Today designers are digitally
developing an architectural vocabulary (most of which has yet to
be constructed) which resembles forms found in nature. Can we
find contemporary ways to “Touch This Earth Lightly”3 through
biomimetic design and the analysis of an animal’s physiological
and behavioral adaptations to local environmental conditions?
Our knowledge of nature should be used more fundamentally as
a resource for sustainable design and for the realization of more
efficient buildings.
Nature offers additional insights for architecture in its ability
to act locally but with an indirect ability to have a global influence. This is illustrated by the interconnectedness of ecosystems.
Ecological landscapes have no fixed boundaries, rather they have
edges that are influenced by surrounding communities. This creates a sequence of locally refined solutions and a network of connections that allow for adaptation to occur over time within each
element of the community or ecosystem.
From buildings to neighborhoods we can design complex ecosystem-like entities, rich with interconnected solutions, which
offer an infinite range of possible results. Ecosystems are rich in
opportunities for allowing each element to coexist with the others. Learning from nature’s efficient interconnectedness allows
designers to consider the potentials for multifunctional responses.
It also offers some examples, as seen in animals, of streamlined
design solutions which might serve as models to ultimately

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reduce and rebalance our footprint on Earth. By embracing bioinspired design processes, opportunities arise to help us develop
man-made environments in harmony with nature, beginning to
eliminate the separation between the built and the living realms.

Owl illustration. Ulisse Aldrovandi, Owl,
Ornithologiae, engraving, 1599. (Image
courtesy of Wikimedia Commons)

Historical Interplay of Bio-Inspired Architecture among
Science, Art and Design   Archeo­logical sites testify to the fact
that nature has been a source of observation and impetus for new
design since ancient human civilization. Examples of Etruscan
artifacts and ancient drawings found in caves demonstrate the
intense need for societies to worship nature. Followers of Greek
polytheistic religions worshipped a constellation of gods and goddesses named after the natural elements. The important Greek
philosopher and polymath Aristotle (384–322 B.C.) put nature at
the center of his scientific studies. In his Historia Animalium he
describes many zoological phenomena. In the ancients’ architecture, nature’s forms are used symbolically and metaphysically.
Nature has always been something to observe, represent, respect,
and worship. However, with the major exception of Leonardo da
Vinci, it was not until the 19th century that thinkers made the
leap from “mere” observation to application. Leonardo could be
considered the first biomimetic designer.
With the Age of Exploration, and increasingly after the discovery of the Americas (1492), an influx of European naturalists
documented their field observations in the form of drawings
replicating nature. As a result, science progressed using tools
borrowed from the art world. History’s greatest naturalists, such
as Leonardo da Vinci (1452–1519), Konrad Gessner (1516-1565),
and Ulisse Aldrovandi (1522–1605), among others, produced
stunning informative drawings. Gessner’s Historiae animalium,
published in Switzerland around 1555, is considered the first
encyclopedic work dedicated to documenting all known animals, particularly through the inclusion of illustrations, drawn
mainly by Lucas Schan. The book generated such an impact
that a few years later all of its illustrations were collected in a
separate book, Icones Animalium (1560). We can attribute the
importance first given to direct personal scientific observations
to Aldrovandi. His watercolor and tempera illustrations, together
with a copious collection of drawings prepared in his studio
by artists hired and supervised by him, depicted real, directly
observed organisms, accurately studied in all their external as
well as internal details.

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In the following century, during the Age of Enlightenment,
1650–1800, scientific explorers and voyagers such as Alexander
von Humbolt (1769–1859) explored the coasts and the inlands of
Central and South America, documenting his discoveries with
beautifully detailed drawings. von Humbolt had a great influence on the explorative work of Charles Darwin, while Ernst von
Haeckel, (1834–1919), biologist and artist, beautifully helped document Darwin’s observations graphically. The intellectual world
became fascinated by the pioneering and explorative findings, so
much so that Denis Diderot (1713–1784), in his Encyclopédie,
dedicated a good 12 volumes to “planches” (plates).

Bat species illustrations. Ernst von
Haeckel, Chiroptera, Kunstformen
der Natur, plate 67, 1904. (Image
courtesy of Wikimedia Commons)

Less tangible and seemingly invisible aspects of nature also
inspired the development of technologies that allowed further
investigation. The invention of the microscope in the late 16th
century in the Netherlands and later the compound microscope
by Galileo Galilei (1564–1642) in 1625 allowed scientists to study
the incredibly close and small as well as the distant using the
same technology. Today, with the aid of the electron microscope,
we can observe the fine structure of a single cell, and with nuclear
magnetic resonance (NMR) spectroscopy, observe protein structures. Robert Hooke (1635–1703) is considered the first to bring
to the public astonishing microscopic images from the invisible to
bare eye from the world of nature.
These early naturalists and thinkers all demonstrated that through
analysis and scientific research, it was possible to create graphical
renditions of the natural world. It was only Leonardo da Vinci
who, contrary to his contemporaries, developed the observations
into design ideas and concepts. Several of da Vinci’s famous
drawings demonstrate the shift from mere inquiries to a world of
human creation and design. In fact, Leonardo may be regarded
as the first biomimetic designer, considering his investigations
on the flight of birds, which developed into the invention of the
first flying machine – the Ornithopter. Another critical da Vinci
contribution is the elaboration of Vitruvio’s de Arquitectura text,
from which he extrapolated and developed the geometric relationships of the human body to the pure geometric forms of the
square and circle. This advanced way of “observing” opened the
world to what we call the “relational aspects.” Leonardo’s drawings provided an essential link to early relational observations in
demonstrating that everything in nature is interconnected, and

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Leonardo da Vinci, Vitruvian Man, pen and ink on paper, c. 1487. (Image
courtesy of Wikimedia Commons)

that clear, relational rules from nature can be applied through
geometry. In this way, creativity comes to play an important role
in the mediation of scientific reason, function, and formal expression through drawing and design.
Concurrent with the great scientific discoveries, art flourished
during the Renaissance. Interestingly, painters also started to
feed their imaginations with naturalists’ observations of landscapes and nature. The art world developed a deep fascination
with the natural world. Artists’ clients, who were often removed
from bucolic landscapes, requested their artists bring those idyllic

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Giuseppe Arcimboldo, Water, oil on limewood, 1563–4. (Image courtesy of
Wikimedia Commons)

views into their homes. While some provided realistic visions of
the natural landscapes, others applied a freer level of interpretation and creativity, distorting and recomposing their visions into a
multiplicity of fantastical variations. Art started to position itself
between the recording of present realities and the imagining of
other places and worlds, thus enhancing spectators’ fantasies and
visions. The creative genius of such inventiveness can be found
in the exceptional work of Giuseppe Arcimboldo (1527–1593).
His fantastic human-like portraits comprised of assemblages of
animals or plant species are exemplary of the inspiration science
and nature provided to art, and how the imagination of the artistic

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mind could recompose such elements in creative and surprising
The 19th century saw a proliferation of engineering and other
applied sciences, with bright minds applying their efforts to the
testing and discovery of many devices. A big shift happened in
1857, when Jean-Marie Le Bris, during one of his long sailing trips
following the flight of an albatross, designed and built the first
bio-inspired flying machine, the “Artificial Albatross.” Another
milestone came with the age of modern robotics in the 1950s. This
time defined a critical threshold of a new branch of engineering in
which scientists were developing bio-inspired devices. Over time
robots have become more and more similar to living organisms.

(Top) Illustration of Artificial Albatross.
Jean-Marie Le Bris, Brevet d’invention,
1857. (Image courtesy of Wikimedia
(Bottom) The living area immersed
in a forest of columns. Alvar Aalto,
Villa Mairea, 1939. (Photograph
courtesy of A. Carr)

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In 20th century architecture there are few relevant case studies
designed by architects informed by nature. Influenced by his mentor, Luis Sullivan (1856–1924), Frank Lloyd Wright (1867–1959)
put at the center of his career his interest in nature. He considered himself the instigator of organic architecture. In his book An
Organic Architecture (1939), he describes how he believed not
only that every building should grow naturally from within its surroundings, but also how the building’s design should be carefully
thought of as if it were a unified organism, in which each element
of it relates to the other, in which each element relates to the other,
similar to ecosystems in nature. Villa Mairea (1939), designed by
the Finnish Alvar Aalto (1898–1976), is one of the finest examples
of nature’s influence in architecture. The forest that surrounds the
house becomes the driving element for the conception of the interiors made of irregular columns and posts. The interiors resemble
the diversity and beauty of nature, from materials to forms, in the
intent to dissolve the separation between the indoor and the outdoor
environments, as in Alto’s words “nature is the symbol of freedom.”
Buckminster Fuller (1895–1983) paid close attention to nature and
its governing systems, as he understood that humans exist in connection with the rest of the living world. Jean Prouve’s (1901–1984)
Maison Tropical (1949) and the later Maison du Sahara (1958) not
only explored ideas of prefabrication and lightness, but are especially important for their design for extreme climates. Frei Otto
(b. 1925), in his prolific career, concentrates on finding the basic
principles of structures in nature. His contribution and influence
are still, today, predominant in the fields of minimal surfaces and
complex geometry. In the late 1950s the Metabolists, a group of

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The Metabolists’ most significant
built contribution that synthesizes
notions of exchangeability and
growth. Kisho Kurokawa, Nakagin
Capsule Tower, 1972. (Photograph
courtesy of M. Nesbit)

Japanese architects led by Kenzo Tange (1913–2005), began looking at cities and buildings as expandable, flexible structures, capable of organic growth and transformation over time. Evidence of
the Metabolists’ notions are significantly demonstrated in Kisho
Kurokawa’s (1934–2007) infamous Nakagin Capsule Tower, which
embodies an explorative and adaptive design attitude for a building
as organism. After decades of postmodernism, attention has been
refocused on nature’s morphologies. Responsibility for this trend
can be attributed to, among others, the work of D’Arcy Wentworth
Thompson (1860–1948). The rediscovery of the 1917 book On
Growth and Form by the mathematician has influenced generations
of architects and designers including organic morpho-architects.
His meticulous work looked at the correlations between biological forms and mechanical phenomena, and his descriptions of the
interrelation between form and growth helped biologists, architects, and engineers find fruitful starting grounds for collaborative
The attention brought to bio-inspired design in the last 15 years has
allowed the emergence of a new group of designers and architects
interested in interpreting, participating in, and collaborating in the
development of architecture which is environmentally and ecologically sound, as well as atmospheric, sensible, and smart. Today
there are many contributors to the advancement of bio-inspired
design and speculative projects within and outside the academic
realm. Only a few examples can be truly considered “biomimetics,”
with even fewer built examples in architecture. Increased attention
to the subject matter has reached a critical mass for further development. Because we are facing some of the most unprecedented environmental challenges of our time, we need architectural responses
that are similarly unprecedented, flexible, adaptive, and performative in function. Biomimetics, by finding direct inspiration from
nature, provides unmatchable lessons to designers and architects.
“A building should be designed so as to minimize the use of new
resources and, at the end of its life, to form the resources for
other architecture.”
- Robert & Brenda Vale
Contemporary Challenges and Interests  Climate change is
arguably the most far-reaching, unprecedented environmental
challenge of our time, and has been recognized as such by the

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United Nations, the national science academies of all major industrial nations, a majority of the world’s governments, and the mainstream public. Although its long-term effects on the environment
are not yet clear to us, it is apparent that the extreme shifts in climate wrought by humans have real repercussions on our ecological equilibrium. The media constantly remind us of the damaging
effects that the perennially melting glaciers, rising sea levels, flooding, drought, fire, heat waves, catastrophic weather events, altered
ecosystems and biodiversity loss, and land and air pollution have
upon our health and economic and political stability.
We are also accelerating the extreme loss of biodiversity through the
loss of habitats caused by deforestation, urbanization, water, land
and air pollution, and other forms of natural resource depletion. The
extinction of species creates large ecological unbalances. When one
species disappears, other species will be affected, provoking a chain
reaction of events. Organisms are facing environmental changes at
a rate that is highly accelerated compared to the normal course of
evolutionary change, and therefore, many species cannot adapt to the
unprecedented ecological pressures that humans have caused. But
what is currently induced is a mass extinction of species, thousands
of which still have yet to be discovered and thousands more classified, jeopardizing entire habitats, the species as indicators of habitat
health, and the services these ecosystems provide to humans. The
International Union for Conservation of Nature (IUCN) estimates
that we are losing species due to anthropogenic pressures at least
1,000 times the natural rate4; 19,265 species of the 59,507 assessed
to date are threatened with extinction.5 We urgently need to change
the way we interface with the natural environment.
As the built environment also significantly contributes to climate
change, it is urgent, and logical, that we consider an integrated
design approach incorporating nature’s time-proven lessons. The
architect’s and designer’s goal is to develop ways to appropriate
and reuse nature’s resources responsibly.
Our challenge today is to balance or eliminate waste by developing ways to build with limited environmental impact and without excessive abuse to the natural world. This applies to how we
design and build, and to how we occupy, maintain, and operate
our buildings. Nature is very complex and highly dependent on
closed-loop cycles. In this homeostatic state, one’s waste is another’s food source. Waste is one of the major causes of resource
depletion and only truly happens when we interrupt natural cycles.

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Bio-inspired design is a transition toward revitalizing, revamping and reinventing our existing environments. This in turn
provides an opportunity for designers of both new and old construction to shift their focus from developing buildings in isolation to considering them as a component of the larger network of
systems intertwined with the multitude of factors making up the
environment as a whole. Performance-based and whole-system
approaches offer infinite possibilities to creatively reinvent,
reprogram, distort, manipulate, rearticulate, and reshape the
environment that surrounds us. Through our vision of a more
compact, more responsible, and more creative way of coexisting, we can come closer to living in a world of equilibrium. The
understanding and design of the building envelope, under particular examination in this book, offers a great opportunity for
addressing such issues.
Performance-based design is necessary in order to lessen our ecological footprint. Innovators are focusing on performative design
techniques used to optimize life-cycle building performance in
an integrated, holistic manner. Integrated systems design coordinates complex systems intended to maximize building functionality while providing for human comfort. The integration of
advanced building systems (e.g., envelope, mechanical, electrical, lighting, and plumbing) combined with sustainable design
practices is our best way forward in this regard. Performative
design can encompass complex geometry, parametric and algorithmic design while moving the process beyond the mere formal
observation of natural forms and patterns. Thus, the goal of this
process is the development of meaningful, flexible and adaptable
relationships between systems from which architecture and its
processes can emerge.
Working holistically does not eliminate the need for step-bystep process-based design, a hierarchical approach to addressing
major topics such as climate and design principles (orientation,
program and space zoning), envelopes and passive strategies (performative and adaptive parametrically complex envelopes, thermodynamic principles, and innovative materials), active systems
(building systems design), and generative systems (renewable
energy sources). Today quantitative and qualitative performancebased analyses offer a comprehensive virtual approach to

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validating the process. Digital tools and assessment methods
allow not only for preventive testing and methodology validation
but also the generation of the building form.
Systems integration starts with the design of the envelope, one of
the most important components of a building, as it incorporates
most of the architectural and building engineering disciplines.
The envelope connects and separates, and acts as a filter between
the exterior world and the internally controlled environment. It
mediates and enhances all relationships between natural elements
and the conventional notion of human comfort. An advanced
understanding of building and environmental design concepts,
principles, and strategies is necessary in order to identify appropriate building systems for different climates and building occupancies. Extensive analysis, testing and systems optimization are
required elements of a performative building.

“I have thought exactly the opposite. Jungles and grasslands are
the logical destinations, and towns and farmland the labyrinths
that people have imposed between them sometime in the past. I
cherish the green enclaves accidentally left behind.”
- E.O. Wilson
Nature and the Built Environment  Presently, there are very
few places in the world occupied “lightly” by man, such as places
where tribal communities remain part of natural ecosystems.
And there are even fewer places left completely untouched by
One can establish a quantitative relationship between the built and
unbuilt environments. However, such quantification alone does
not necessarily lead to a definitive answer we can implement to
produce real change. We must consider a qualitative assessment
of this quantitative relationship if we are going to productively
shift and reverse the currently large human ecological footprint.
A constructive attitude toward conservation and restoration policies starts with the healing of sites’ ecological and architectural
systems at the master planning scale. We must modify the way
we design our buildings so that they will not contribute to the

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depletion of natural resources, but rather work to reintroduce
resources back into ecosystems, ultimately benefitting the environment at large.
An example of a restorative approach to design can be found in
the project “A Model Community at Salton Sea” by im studio mi/
la and collaborators. The traditional model of growth is a zero
sum game with discrete land use typologies, where growth in one
area can only happen at the expense of another. This project provides a paradigm shift in accommodating growth – by capturing and integrating systems that are cyclical in nature and time,
with each cycle rejuvenating and healing the surrounding ecology
rather than eroding it. The approach is holistic, in that it considers
interrelationships between all processes fundamental to sustaining life and preserving nature – water and energy cycles, agriculture and seasonality, production and the exchange economy, as
well as the social needs of a multigenerational community. The
applied strategy, however, is hinged on the notion of restoring
scarred landscapes, making them givers of life, and enhancing
their integration into the surrounding ecology.

Scarred landscape and debris of
dead fish off the shores of the Salton
Sea, CA. (Photograph courtesy of R.

As a result of untrammeled population growth, the development
of cities and degradation of rural sites have confined nature to an
increasingly smaller and limited area intended for conservation,
monitoring, and biodiversity preservation – both outside of and
within small and large cities. The concept of nature in urbanized areas has transformed and redefined our understanding of
what nature is and how to interact with it. The question is if citydwellers still maintain a relationship with nature in their urban
lives. And if the sense of beauty we associate with nature is in
contemporary living something we relate to only in a romantic,
bucolic, and melancholic way, rather than in an interactive, daily,
practical way.
Nature is dynamic and adaptive, thus allowing for restoration.
Healing degraded areas is the first step toward a healthier relationship between nature and the urban environment. The second
step lies in architecture’s long- and short-term response to nature’s
ever-changing conditions; architecture needs a new context in
terms of environmental factors that define a place, beyond the
traditional contexts of physical surroundings, geometries, architectural styles, and stylistic traditions.

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Master plan depicting inputs on the left, and the outputs produced by the
community filtered through the system on the right. Model Community at
Salton Sea, CA, 2006. (Image courtesy of im studio mi/la and collaborators)

The notion of aesthetics also plays a substantial role in incorporating nature into the built environment, as it establishes emotional
attachments and garners respect. In human society, beautiful
places tend to be valued more. By designing aesthetically pleasing things, we can help their conservation. Issues of aesthetics
in architecture have been discussed since the time of the ancient
Greeks, a society highly interested in such philosophical aspects
of life. In science, beauty is also often present but discussed in
different terms, such as through laws and mathematical formulae. The semiotician Umberto Eco, in his book History of Beauty,
offers a peculiar reading of how this concept often relates to the
feminine attributes of curves and grace. But architects and scientists seem to disagree on the importance of simplicity versus
complexity, elegance, and symmetry as elements that would conclusively define what is pleasing to the eye. Neither architecture
nor science can agree upon which attributes constitute beauty,
as there is no universal definition of it. Biomimicry might seem
to embody objective aesthetic notions because humans are hardwired to find nature attractive. In bio-inspired design, beauty is
something that is cohesive and respectful of others’ surroundings.
This is because nature provides numerous attractive examples,
at all scales. In fact, from the invisible to the visible, nature

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seamlessly integrates all the parts of a whole, with not only her
forms but also through functions, processes, and systems.
“When I’m working on a problem, I never think about beauty. I
think only how to solve the problem. But when I have finished, if
the solution is not beautiful, I know it is wrong.”
- R. Buckminster Fuller
Novel Practices in the Built Environment: Dynamic,
Atmospheric and Active  Is it possible to create an architecture
that is responsive and flexible, engaged and adaptive to an equally
dynamic environment in which it is integrated, not simply in a
stylistic way, but rather in an active, participatory, and even regenerative way? Can we transform our cities from the inflexibility of
a few iconic buildings surrounded by nondescript urban fabric
into places made of elements that participate in the nourishment
and well-being of their inhabitants? Can we avoid the seldom
attractive and instead embrace the meaningfully beautiful that
will add value to our everyday and work lives and enhance our
quality of life?
Despite the countless number of design competitions and investigations (particularly on paper) that have actively responded to
environmental stimuli and cues in the last few decades, there are
still a limited number of built examples that move us toward a
vision of a dynamic, responsive, and sensory charged architecture. Because it is not yet mainstream, adaptive architecture is
very hard for the general public and clients to imagine, further
limiting its acceptance and proliferation.
Since the 20th century many architects have been intrigued by
and have attempted to build environmentally responsive buildings. Yet only a few succeeded. The most recent and perhaps
most renowned is Jean Nouvel’s Institute du Monde Arab in Paris
(1987). The building’s south facade is articulated by a continuous Moresque-inspired screen which blurs the reading of the elements, changing the perception of what is typically understood
as window and wall, making them indistinguishable and interchangeable. Computer programmed, mechanically activated
“irises” respond to the varying sunlight intensity by opening or
closing, thus optimizing interior daylight and comfort. From the
street the building reads at times as a whole wall and at other

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South facade. Jean Nouvel’s Institut du Monde Arabe, Paris, 1987.
(Photograph courtesy of J.C. Martin)

Iris detail of a single operable panel
for the Institut du Monde Arabe.
(Photograph courtesy of J.C. Martin)

times as a whole window. This ambiguous perception, along with
the technology that allowed the mechanisms to operate the oculars, was a breakthrough in the history of performative architecture. Drastically different from a typical curtain wall façade in
which each individual element would aspire to be all window,
all transparent, the library screen south façade appears as an
uncanny object in the cityscape, acting and reacting to changing lighting conditions by constricting and dilating – analogous
to the human iris. The screens of the facade stand proud against
the Paris skyline, yet today, most of the mechanical “pupils” are
nonfunctioning. While the beauty of the building remains almost
intact, its usability and kinetic aspects are now lost. This building
has taught that both the engineering and architecture of highly
mechanized buildings must be designed to last. Both, in fact, have
to deal with the reality of the complex movement while the technology must be developed with long-term ease of maintenance
and durability in mind.
The operability of a building may well be the main challenge in
the architectural design process today. In fact, the mechanisms
must be designed to match the life span of a building. Ultimately,
the engineering of highly adaptive building systems determines

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their success or failure. Largely, at this point in time, experimentation in the area of active façade technology is often realized only
in temporary installations. Illustrious case studies of such ephemeral experiments, such as the Crystal Palace by Joseph Paxton
(Great Exhibition of 1851), the Blur Building by Diller Scofidio
+ Renfro (Swiss Expo 2002), the various pavilions of London’s
Serpentine gallery, and New York’s PS1 summer installations,
showcase the most avant garde designs and tests of movement,
structural reaction, and adaptation to environmental conditions
encountered in transient expositions.

Olafur Eliasson, “The Weather
Project,” Turbine Hall, Tate Modern,
London, 2003. (Photograph courtesy
of D. Thair)

Art has helped advance architecture’s investigations and suggests
many possible ways to move forward, away from the static and
mute massing of the current built environment to more active,
responsive, atmospheric and responsible practice. As early
as the 1970s, the work of artists such as Gordon Matta-Clark,
James Turrell, Dan Flavin, Dan Graham, and Robert Smithson
has provoked our spatial imaginations with its complex interplay between architecture and light, sky and atmosphere. One
of the major exponents of such artistic tradition today is Olafur
Eliasson. With his work he helps us to experience space and time
In his dynamic installation “The Weather Project” in London
(2003), he created an artificial climate within the Turbine Hall of
the Tate Modern. Throughout the day, mist was released indoors
under the heat of a large artificial sun made of hundreds of lamps
radiating yellow light. While lying down on the floor the visitors could see their shadows mirrored as tiny black dots in the
hall’s ceiling. The transfer of atmospheric effects into an interior
space provided the spectators with a sensory experience akin to
the light and heat that rises every morning and sets every evening, giving rhythm to our lives. With “Your Atmospheric Colour
Atlas” shown at the 21st Century Museum of Contemporary Art,
in Kanazawa Japan (2009–2010), Eliasson engaged the viewer in
an immersive colored environment saturated with fog, forcing the
viewer to have to negotiate between actual and perceived spatial
realities, as they lost sight of the edges which define the space. By
heightening the visitor’s awareness of perception, the augmented
reality typical of Eliasson’s work provides a powerful moment of
clarity. The profundity of such phenomena eludes wordy descriptions, registering instead more sharply in direct experience.

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Interior rendering for the proposal for the Museum of Contemporary Art in
Wroclaw, Poland, 2008. (Image courtesy of Philippe Rahm Architects)

Illustrations generated by climatic
software used to model the various
climates of the museum to appropriately organize the program. (Image
courtesy of Philippe Rahm Architects)

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In architecture, some of the most sophisticated phenomenological explorations are found in the work of Philippe Rahm. His
interest in affect and performance explores the sensation versus
the image, while focusing on meteorological principles to control the interior environment. Rahm’s predilection is, in fact, to
use gradients of heat and humidity to define space. His entry for
the Museum of Contemporary Art in Wroclaw, Poland (2008),
combines a deep meteorological and atmospheric understanding of the museum as both a container protecting the arts and a
place for contemplation, visitation, and experiential exploration.
The program is organized by carefully studying temperature in
air stratification: within the cooler environments the majority
of the square footage is dedicated to areas of less occupancy
(16°C), while the galleries and offices (22°C) are located at the
increasingly warmer levels. Driven by climatic requirements the
architectural volume emerges in the form of a stepping sectional
diagram. The architect’s interest in the ephemerality of air and
in its “soft solid”6 state has led him on an investigation focused
on “ecology (author’s note: rather than program in terms of area
and volume) as a driving tool for changing the design process. A
process for inventing a new way of living.”7

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An external rendering of one of the galleries for the Yeosu Oceanic Pavilion.
Tom Wiscombe in collaboration with Roland Snooks — Yeosu, South Korea,
2010. (Image courtesy of Tom Wiscombe)

R&Sie’s work is singular through their use, manipulation, and distortion of environmental conditions, and construction of dynamic
architectures, which result in their mechanistic expression. The
team overexposes and almost fetishizes the use of ecological systems to enrich and inform their architectural projects. A significant built example is the house enclosure constructed in 2008, I’m
Lost in Paris, where the inhabitant assists in the explicit use of
the exhibition and the changes occurring in the environment. The
hydroponics system nourishes ferns with a bacterial concoction,
while the hundreds of glass beakers in which they are contained
provide light for the interior spaces. The resultant “living facade”
indexes growth, recording environmental changes.
The work of Tom Wiscombe is a passionate exploration of morphology, as relating to the original meaning of the word: the
study of natural forms and structures together with a fascination
for composite materials. “Ultimately, multi-materiality allows
for variable opacity, color and depth effects never before seen
in architecture, and possibly only in the transparent head of the
Pacific barrel eye fish or the deep colorful organs nested inside
the translucent body of the Costa Rican glass frog.”8 Tom’s rigorous morphological studies on nature’s emergent behaviors find

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Interior view of Hylozoic Ground. Philip Beesley, Canadian Pavilion at the
Venice Biennale, 2010. (Photograph courtesy of I. Mazzoleni)

their strength in his interdisciplinary collaborative process with
biologists, structural engineers, and computational architects.
The Yeosu Oceanic Pavilion (2010), done in collaboration with
Roland Snooks, exemplifies a new phase of exploration in which
the use color as a critical element of communication, beyond
its narrow indexical association, is at the center of Wiscombe’s

Detail of the flexible meshwork for
Hylozoic Ground. (Photograph courtesy of I. Mazzoleni)

Philip Beesley’s work is characterized by his interest in lightweight textile structures that incorporate living organisms. In
Hylozoic Ground, the installation for the Canadian Pavilion at
the Venice Biennale (2010), he explores the use of synthetic protocells, a form of biology that can transform and interact with
both the presence of people and the surrounding environmental
conditions. This material investigation is often complemented
by computational studies in mirroring the way genetics develops
its understanding of living organisms, resulting in an immersive
Architects today are also exploring possible future applications
of high performance materials capable of self-healing, growing, and decomposing as living organisms. Biomaterials are

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being developed for biomedicine and defense applications and
are inspiring many architects. The most important next steps in
architectural advancements will be found in nanotechnologies,
materiomics,9 metamaterials, and biosmart materials’ ability
to change properties (thermal, luminous, acoustic) or exchange
energy (phase change) depending on external environmental
Materials science, together with emergent technologies, computational and parametric design, find common ground in the complexity of nature and the processes governing it. Nature’s fundamental
mathematical laws are parametricized with the intention to create
performative yet surprisingly bottom-up novel forms.
The iterative nature of methodologies associated with scripting, or
coding, often used as form finding, can be successfully applied to
any ordering system, including performance-based design. Some
professionals prefer to use it in its pure state, algorithmic and parametric only, whereas others use it as the first generative step, to then
corrupt it, injecting chaotic and messy elements in the subsequent
iterations to provoke distortions, “polluting” the original pureness of the mathematical matrix. In either case, the final moment,
the “done” moment, is determined by human determination and
likeness of the seen result. While the computational process provides the starting point, only rarely it determines its conclusion.
In performance-based design the optimization of energy resources,
the limitation of energy consumption, and the optimization of passive systems help shape the end product.
Learning to be attuned to environmental conditions around us,
as organisms have adapted to their local environment, is the most
deep-rooted lesson we can learn from nature and is the book’s
inspiration. Animals have anatomically, physiologically and
behaviorally adapted to their environment for survival. Our species, with our great intellect, has found ways to bypass the biological course of evolution with buildings, falsely thinking that we can
control this divergence and procure only the benefits. While this
venture has often failed, bio-inspired design can help us integrate
once again with our evolutionary paths. The book focuses on a
select number of lessons from nature and suggests some possible
solutions. Respectful and responsible steps in design are needed to
induce action and change in existing highly anthropomorphized
environments without disregarding, but rather regenerating, their

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present conditions. The currently built environment should be
seen as something in need of healing. Priority should be given
to restoration and transformation, rather than continuing to irresponsibly expand into more — and therefore fewer — rural areas.
Nature provides an infinite source of precedents that have evolved
through the years to provide optimal solutions to innumerable
challenges of life and survival. Our senses perceive nature’s efficient solutions as beauty and our minds are inspired by these
elegant natural systems. The analytical brain seeks to understand
nature’s driving laws and principles, and the creative brain is aesthetically stimulated by these biological creations. Nature thus
serves as an endless source of inspiration for those in search of
novelty and beauty in efficient and effective design. The transition
from bio-inspiration to biomimicry depends on both analytical
and creative talents combined with expert knowledge in a conscious effort to synthesize innovative solutions through a persistent search of nature’s repertoire.

Urban (Dis)solution, charcoal on
Strathmore, 2009. (Image courtesy of
im studio mi/la with D. Kim)

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“Nothing in biology makes sense except in the light of evolution.”
- Theodosius Dobzhansky

How Biology Informs Architecture
Biology is the study of life. Defining life is challenging, and there
is no absolute consensus for its definition. Most scientists agree,
however, that several traits characterize all living things. Living
organisms undergo chemical processes for energy and maintenance. Cells gain energy through a constant cycle of breaking
down matter and constructing matter into cellular components.
Living beings reproduce, either sexually or asexually. They are
able to evolve and adapt to their environment over time, so that
characteristics of a species change from generation to generation.
They respond to stimuli, and in doing so, they maintain homeostasis. Life is dynamic, meaning that all traits that characterize
living beings involve dynamic processes.
The basic unit of life is the cell; it is the building block of all life
forms. A living organism can consist of only one cell or of multiple differentiated cells. Though single-celled organisms, like
most bacteria, seem very different from organisms with millions
of cells, they are still connected, as each undergoes all of the basic
processes of life.
The field of biology seeks to understand the patterns and processes that govern all living things. Biological patterns and processes are incredibly diverse and complex, and many disciplines
of biology exist to understand how life works. For example,
biochemistry is the study of chemical processes fundamental
to living organisms such as metabolism and cell signaling. The
study of the components of cells, how they work, and how they
interact with their environment is the focus of cell biology. The
discipline of genetics is concerned with gene function and how
genetic information gets passed from one generation to the next.
On a larger scale, evolutionary biology seeks to understand how
species originate and how they change over time, while ecology
is concerned with broad scale patterns of interactions among
organisms and interactions among organisms and their environment. Biology is also interdisciplinary; it integrates with other
fields. Biomathematics uses models to understand and predict life

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Diagram of a generalized animal cell. (Image courtesy of im studio mi/la)

processes. Bioinformatics uses information technology to store
and analyze biological data. Bioengineering uses biological principles for application in design and manufacturing.
Scientists employ a specific methodology, called the scientific
method, to acquire knowledge about the natural world. This
methodology is the basis for all modern scientific research, as it
is evidence based and strives to be objective. The process involves
inductive reasoning, moving from observations of the world to
deriving general patterns and principles about how the world
works. First, observations are made of a phenomenon in the natural world. Then, a hypothesis is formulated to explain those observations. Predictions are made that follow from the hypothesis. In
science a hypothesis must be testable. It needs to be framed so that
some kind of measurements can be taken in order to determine
that the hypothesis is true. A hypothesis must also be falsifiable;
scientists must be able to show that it is incorrect. Consequently,

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an observational or experimental study can be designed with the
goal of testing the hypothesis, and the results from the study are
used to make an interpretation of the hypothesis. The data will
either support or reject the hypothesis. Through multiple, sequential testing of related hypotheses scientific knowledge is gained.
The scientific method is necessarily iterative. An idea has to be
tested over and over again before it is accepted, and even then,
new evidence may come to light that refutes previously supported
hypotheses. Therefore, ideas in science are always changing. The
process of conducting science is a repeatable, ever-refinable one,
where hypotheses are constantly tested and nothing is ever proven
In contrast, the design or creative process differs from the scientific method and is more a deductive reasoning process, beginning
with the knowledge and study of general principles which in turn
lead to specific solutions. Building design emerges in response to
the rise of contextual questions and the necessity for a specific
population to provide for needs related mainly to function, protection and comfort.
The projects go through several steps, from the conceptual to
the detail of building components. While the conceptual and
schematic phases are characterized by their intuitive, open, even
“messy” process, decisions become more definitive with each
iterative step and less able to change as the design approaches
the construction phase. This contrasts with the scientific method,
which allows for continuous change to be taken into consideration
and tested. The early creative process, especially in its beginning
phase, is of specific interest in this book. Architecture looks for
precedence to confirm, validate, but most importantly expand, its
design exploration in search of innovative ideas. Nature itself is
offering us a meaningful source of new precedents for design.
Humans are understandably driven to explore the world around
us, and for good reason. Throughout the evolutionary history of
our own species, biological knowledge has been imperative for
survival. Humans needed to learn which plants were safe to eat
and which ones were poisonous. It was crucial to understand the
behavioral patterns of predators to avoid becoming prey. In one
form or another humans have gained biological knowledge over
millennia. However, despite the number of biological disciplines
and the amount of time we have been studying natural phenomena,

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we still know relatively little about the natural world. There are
vast numbers of species, biological processes and biological interactions that are yet to be understood. We do not completely understand how life evolved, how many species exist on Earth, or the
details of how the genetic code operates. These are just a few of
the fundamental questions that biology still has to answer.
The disciplines of design and architecture can extrapolate knowledge from the biological world in order to improve the way
humans live. The field of biomimicry applies biological principles to design in two main ways. First, a focus of biomimicry is
to understand the dynamic context within which we operate and
place buildings into the environment. Study of the natural world
takes into consideration the interconnectedness and diversity of
nature, and architects can learn from this perspective in order to
design elements that can become integral parts of natural systems. Biomimetic designers strive to be more considerate of the
environment, to be less invasive and more conscious about the
fact that humans live among other components of nature. Second,
designers can learn about the functional aspects, or adaptations,
of organisms and translate those principles into design. Biological
form is studied for its function, which is a basic precept that can
be imbued into architectural design through the study of precedents. The starting point for such investigation can occur at either
large design scale, such as that of a city or neighborhood, looking
for inspiration in the complexity and interrelatedness of ecosystems, or by choosing one element of an animal as focus of inspiration, for example, starting with an animal skin to use as model for
the design of a building envelope. In this book we develop both
principles, recognizing that nature teaches us that everything is
connected; therefore, our study of the building envelope is not
meant as design of a part in isolation, but rather the opposite,
which focuses on that part in the context of the whole.
Evolving and Adapting to Survive  Life on Earth has changed
over time. Living beings appeared approximately 3.5 billion years
ago in the form of single-celled, aquatic relatives of bacteria. Since
then, millions of species have come into existence and have taken
myriad forms. This process is called evolution and is the unifying theory of biology. Evolution is the conceptual framework that
brings together all the disciplines of biology, because it explains
how living things came to be, how they are related to each other,
and how they function. In its most basic form, evolution simply

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external skeleton



nervous & vascular system

An evolutionary tree, or phylogeny, of animals. Major evolutionary adaptations
are depicted on the tree. (Image courtesy of im studio mi/la)

means change in the characteristics of a population over time.
For organisms with short generation times, this process is directly
observable. All living beings share the same genetic code, and
evolution occurs when the frequencies of different forms of genes
change over time.
Four major mechanisms of evolution are recognized: mutation,
natural selection, genetic drift, and gene flow. Mutation is the
basis for all genetic variation. It is the spontaneous change of
genetic material, which can be caused by external factors such
as radiation or chemicals, or by internal, genetic processes, such
as errors that occur when DNA is replicated. Genetic variation is
the underpinning for all other processes of evolution. It must be
present in order for evolution to occur.
The mechanism of natural selection was a revolutionary theory
proposed by the English naturalist Charles Darwin (1809–1882),
and it explains how organisms adapt to their environment. The
theory of natural selection states that individuals within a population have variation in their traits. Because of the inherent
variation in populations, some individuals will have traits that
are better suited to the environment than others. Individuals

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possessing those traits will survive and reproduce more than
others in the population. If the traits are heritable, that is, if the
traits have a genetic basis, then this process leads to change in
the characteristics of populations over generations because individuals with the most well-adapted traits pass them on to their
offspring. An important prediction of this process is that the
traits being selected for are only those that allow organisms to
survive in the environment that they live in at the time. When
the environment changes, which it does continuously, the traits
that were previously successful may not be adapted to the new
environment. If there is enough variation, the characteristics of
the population may change again. If not, then the population may
become extinct.
The other two mechanisms of evolution do not actively help populations adapt to their environment, but do change the frequencies
of genes in a population. Genetic drift occurs when gene frequencies change due to random sampling. The frequency of traits in
a population that do not necessarily have an adaptive value can
vary over time based on which individuals happen to mate and
pass on traits to the next generation. Gene flow occurs when individuals from one population migrate to other populations. This
migration tends to add genetic variation into a population as well
as alter the frequencies of forms of a gene in populations.
Biologists recognize two major scales of evolution, which in
reality are part of the same continuum. Macroevolution refers to
processes at or above the species level. Studying speciation, the
process whereby new species evolve, or how traits of organisms
have changed over millennia using the fossil record are examples
of macroevolutionary research. Most people think of evolution at
this scale. However, evolution can occur over smaller time frames
as well. Evolution within a population or a species is referred to
as microevolution. The frequencies of forms of a gene are usually
constantly changing within populations, even though they may
not yield large scale, obvious changes and do not give rise to new
species. The development of antibiotic resistance in bacteria is a
prime example of microevolution.
On the macroevolutionary scale, key innovations have led to the
success of many groups of animals. Key innovations are traits or
characteristics of a group of organisms that allow it to diversify
and give rise to many species. These traits may give organisms

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the ability to acquire more resources or they may allow them to
exploit previously unavailable resources, promoting expansion
into new habitats or niches. There are many examples of key
innovations, and they range in scale from the evolution of wings
and flight to the evolution of cusps on teeth that allow for the use
of new dietary resources. The evolution of the eye exemplifies
a particularly successful key innovation, because so many more
animals possess eyes than do not possess them.
The evolutionary interplay between predators and prey is often
interpreted in terms of coevolution. Coevolution is defined as
reciprocal evolutionary change between species; an evolutionary change in one species causes one in another species. Animals
do not exist in a vacuum. They interact with many other organisms in their ecosystem, and these work to influence evolution in
each other. Besides the coevolution between predators and prey,
another classic coevolutionary example is between parasites and
their hosts. Coevolution is an important aspect of evolutionary
study given that it is likely that every living species on the planet
has coevolved with other species.

(Top) The origin of the eye is thought
to be a key innovation in the evolution of animals. (Photograph courtesy of N. Amin)
(Bottom) Coevolution is common
between plants and their pollinators,
such as bees. The plants have their
pollen spread to other plants, and,
in turn, pollinators get rewards such
as nectar. (Photograph courtesy of S.

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The concept of adaptation to the environment has not traditionally been central to architectural design, yet to improve our ecological footprint it needs to be taken into consideration. The pace
of growth of human building construction worldwide is more
rapid than the rate of evolution operating on biological time­
scales. Having more buildings equates to less animal habitat, less
resources and more pollution. Slowing the pace of building by
humans while being cognizant of how architecture can fit into the
environment around it will help achieve the goal of lessening and
perhaps improving our negative impact on the Earth.
Climate and Biomes  Throughout our history humans have
caused major, negative impacts to the planet. As technology has
progressed and our population has grown, the rate of humaninduced changes has rapidly increased. Much of what has driven
these negative effects on the environment is the absence of being
willing to change even given the harm we know we are causing.
A major issue facing the planet now is global climate change. The
increase of carbon dioxide, primarily due to the burning of fossil
fuels, together with the release of other greenhouse gases into the
atmosphere, is having significant impacts on regional and global
climate patterns.

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Because of the spherical shape of the Earth the sun’s rays are not equally
spread across the planet. Areas around the equator receive the most direct
light, which is why they are warm. In contrast, areas at higher latitudes are
cooler, because radiation from the sun is spread over a larger area. (Image
courtesy of im studio mi/la)

Each terrestrial region in the world is characterized by major vegetation types called biomes, and each biome is associated with a
particular set of climatic conditions. The term “climate” is used
to refer to a suite of variables such as temperature, moisture, sunlight and wind that characterize a region when considered over
long periods of time. Weather is also sometimes used to refer
to the same set of variables, but over much shorter time scales.
Large-scale climate patterns are regulated by the Earth’s shape,
tilt on its axis, and orbit around the sun. Global temperature patterns are governed by the angle at which the sun’s rays hit different parts of the Earth. The spherical shape of the Earth means that
the equator receives more direct solar radiation per unit area than
regions closer to the poles; therefore, equatorial regions are warm.
Higher latitudes are cooler, because the same amount of sunlight
is spread over a larger area and has traveled farther to reach the
Earth’s surface.
Global patterns of precipitation are influenced by cycles of air
circulation. Areas along the equator receive the most moisture,
while areas at around 30° latitude north and south are the driest
on Earth. This is due to the differing densities of warm and cold
air. Warm air is less dense than cold air, causing warm air to rise.

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90 N
North Pole

60 N

30 N

0 N

30 S

60 S

90 S
South Pole

A major driver of global climate patterns is the way in which air circulates
latitudinally. Cycles of evaporation and condensation create areas with moisture around the equator and at 60o latitude, while latitudes of 30° and 90°
are dry. (Image courtesy of im studio mi/la)

When the sunlight hits the equator it causes warming of the air as
well as evaporation of water. This warm, moisture-laden air rises,
cools and condenses in the atmosphere, and then falls as precipitation. Because the Earth spins on its axis, rotating air currents
move air in a north to south pattern, creating cycles of evaporation,
condensation and precipitation. As the warm air moves toward
the poles it cools, and at around 30° latitude, begins to descend.
This cool dry air mass absorbs moisture from the Earth’s surface,
creating arid conditions at that latitude. At latitudes around 60°
the cycle repeats; air rises, cools and releases precipitation. The
cold, dry air then travels toward the poles, where it absorbs moisture and creates cold, dry tundra.
Seasons, annual fluctuations in temperature and precipitation, are
caused by the Earth’s 23.5° tilt on its axis. Summer occurs when
either the northern or southern hemisphere is most tilted toward
the sun as the Earth circles around it, and winter occurs when
each hemisphere is tilted away from the sun. Regions near the
equator do not experience major changes in temperature, but precipitation does vary.

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cold temperate



dry / temperate

The Köppen classification system, one of the most widely used systems,
divides global climate into five major types. (Image courtesy of im studio

Ocean currents, which determine water temperature and greatly
affect air temperature, are influenced by air circulation patterns,
the rotation of the Earth, and the intensity of sunlight hitting the
Earth’s surface. Warm waters at the equator are carried toward
the polar regions along the eastern coasts of the continents, while
cold water traveling toward the equator moves along the west
coast of continents. These patterns lead to differences in the distribution of warm and cold water in the oceans, as well as changes
in the distribution of warm and cold air on the Earth’s surface as
it travels with the water.
The most common climate classification scheme is called the
Köppen system, which defines five broad climate types: tropical, dry, highlands, cold temperate, and polar. These types can be
further divided into secondary classifications. These secondary
classifications often correspond to major biomes, depending on
classification scheme.
Biomes tend to have similar communities of animals within
them, because the animals have evolved in similar climatic conditions. Climate, however, has changed over time, and accordingly the distribution of animals on Earth has changed with it.

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The field of biogeography focuses on understanding the distribution of species in space and time. The location of species
is a result of current and past climate. The Earth’s climate has
changed over millennia due to slight but continuous changes in
the shape of the orbit of the Earth around the sun, the tilt on its
axis, and in the orientation within its orbit. Over geologic time
the amount of sunlight hitting the surface of the Earth and how
air and water circulate are altered. Subsequently, warm periods
that can last for thousands of years alternate with cool periods,
often accompanied by glaciation. Additionally, the movement of
the continents has shifted significantly over time, which causes
changes in biogeographic patterns. The Earth’s crust is divided
into large, thick (50–250 miles) plates that constantly move over
the soft mantle underneath. It has been hypothesized that a giant
supercontinent existed about 200 million years ago that broke
up over time, leading to the current position of continents. This
process of continental drift explains many patterns in plant and
animal communities over the Earth. Closely related organisms
can occur in very different places if they evolved on a land mass
that later split apart. Conversely, unrelated animals can have
similar sets of adaptations if they have evolved in similar climatic regimes.
Ecosystems and Biodiversity  Over the last 3.5 billion years, a
remarkable number of species evolved. Almost two million species have been named, and it is likely that millions more have yet
to be discovered. Biodiversity, or the variation of all life forms,
can represent different levels of biological organization. Genetic,
ecosystem and species diversity are all considered part of biodiversity. Most commonly, though, biodiversity refers to the number and composition of species in a given place, and maintaining
biodiversity is incredibly important for ecosystem stability and
function. Ecosystems are all the co-occurring organisms and
abiotic conditions in a particular area, and they function as an
integrated unit. Many ecosystems occur within a biome. Some
ecosystems have incredibly high levels of biodiversity, such as
rainforests, while others, such as the tundra, have sparse numbers
of species.
Estimates of undiscovered species range from five million to
fifty million. In general, species that are large and charismatic
have been named and studied significantly more than small or
microscopic species, such as invertebrates, fungi, and bacteria.

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A pie chart representing estimated species richness for major groups of
organisms. Data from Purvis and Hector (2000). Image adapted by im studio
mi/la. (Image data courtesy of Nature Publishing).

Approximately 15,000 unknown species are identified each year,
most of them being from less well-known and studied groups.
Though most people think of plants and animals when they think
of biodiversity, the greatest amount of diversity is composed of
single-celled organisms. These organisms are incredibly diverse
in how they live. Some use oxygen for respiration, some convert
light energy to carbon compounds, and some even use inorganic
chemical compounds for energy.
Because they function as a unit, the processes that govern ecosystems include both random, or stochastic, events such as natural
disasters, as well as the more predictable interactions and feedback among organisms, such as predation, competition, parasitism, and mutualism. The outcome of interactions among species
can be destructive to both interacting organisms, as in the case of
competitive interactions for resources. Alternatively, the outcome
may be positive, as in a mutualism, which is a symbiosis that benefits individuals of both interacting species. Finally, an interaction
can have a negative impact on one species but a positive impact on
another, as in the case of predation where one organism is eaten
but the other gets nutrition to sustain life.

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Neotropical rainforest ecosystems, such as this lowland forest in Ecuador,
hold extremely high levels of biodiversity. (Photograph courtesy of S. Price)

The loss of biodiversity and ecosystem function induced by
human activities is occurring at an alarming rate. Dozens of species are likely going extinct each day, many without ever having been discovered. It has been estimated that current extinction
rates are 100 to 1000 times greater than baseline level extinction
rates over geologic time. This current level of extinction is comparable to those documented during mass extinction events, such
as the Cretaceous-Tertiary extinction event that killed the dinosaurs, during which 75% or more of species become extinct over a
short period of geological time. There are many causes of species
loss; chief among them are habitat destruction, climate change
and the invasion of non-native species.
Ecosystems provide many services and resources for humans, and
the loss of species within them can have negative impacts for the
human population. We extract wood and food from ecosystems,
we rely on them to recycle nutrients, plants absorb the carbon
dioxide that humans produce, and they help regulate climate.
Conservation biologists Paul and Anne Erlich have warned that
the health of ecosystems depends on all of the biological diversity
within them; the removal of any one species has some measurable, negative impact on the system and the removal of more and

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more species will at some point break the system. Consequently,
ecosystem function will be lost. In the face of human-mediated
environmental change, conserving higher levels of biodiversity
can provide greater resilience against ecosystem collapse and
mass extinction.
Biomimicry seeks to link complexity, biodiversity and coexistence, using them as precedents for principles of design where,
even when the starting point of the design investigation involves a
singular element, its development is understood as part of a whole
system that interrelates natural and man-made components,
allowing for a beneficial coexistence.

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“Humans have a tendency to fall prey to the illusion that their
economy is at the very center of the universe, forgetting that the
biosphere is what ultimately sustains all systems, both man-made
and natural. In this sense, ‘environmental issues’ are not about
saving the planet—it will always survive and evolve with new
combinations of atom (sic)—but about the prosperous development of our own species.”
- Carl Folke

Lessons Learned: Biology to the Built Environment
All humans have an inherent connection to nature on some
level. Our innate ideas of beauty are connected to forms in the
natural world. Humans feel an instinctive, primal draw to nature
because we have been very closely connected to it for the entire
time we have existed on Earth. The desire to connect to nature
is ever-present, and this longing provides an entry point for
Recent advances in science and technology allow for more elements of the natural world to be used as inspiration. There are
so many facets of the living world to study – from ecosystems
to microscopic organisms and even DNA. There are big pushes
to advance scientific innovation and creativity as well as the globalization of information. Information can be transferred across
the world almost instantaneously, and interdisciplinary work is
facilitated by these advancements.
Architecture has a lot to learn from science, both from a methodological perspective, as a source of design precedents, and as
a springboard for new ideas. As new research is developed in
the fundamental as well as applied sciences, architecture can
increasingly find models for bio-inspired design through applied
concepts and the discovery of new study tools. For example, the
materials sciences are entering an exciting new phase related to
multi-scale material systems – materiomics – which looks at not
only the incredibly small, but also at how observations can be
made in an interrelated way across different scales.
The method of investigation presented in Part II starts with one
element of biology, the skin, and one element of architecture,

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the building envelope, yet these components also serve as larger
ideas, as bridges between realms. The skin and building envelopes exemplify the dialogue between internal and external
worlds as they respond to inputs from both. They protect us from
the environment while keeping us connected to it. The skin is just
one of innumerable aspects of biology that can serve as inspiration for design, but the use of skin and building envelopes here
clearly defined a starting point for our investigation and vision.

Polar bear, Ursus maritimus.
(Photograph courtesy of M. Johnson)

Our exploration of skins and building envelopes unfolds through
a marriage of methodologies – combining the linear and analytical scientific method with the more lateral, synthetic, deductive
creative design process. The difference between these processes
enhances the potential for design. Scientific ideas are never certain and can never be proved absolutely. The search for more and
more support for scientific hypotheses is never-ending. When
thinking about bio-inspired design, we need to consider how its
success is measured. In science, if a hypothesis is shown to be
incorrect, it is discarded. In design, the measure of success has
to do with whether the idea is a springboard for further inspiration and for what emerges in the design realm. In design the
designer defines the goals. If the goals are met, the design is successful. In science the objective is to understand nature to its fullest extent. In architecture, nature is used to inspire design and
its processes whether or not the underlying theories about nature
remain valid through time. If the resulting design concepts that
come from biological observation in architecture are innovative
and move design thought forward, it does not matter if the science
is later proven wrong. The determinant of the project’s success is
its function. In this book, we offer two case studies (in Part II),
each with a particular story.
The polar bear project, the first animal studied for this book, was
developed prior to the publication of a paper correcting a previous
understanding of how this animal’s hollow hair follicles might
be contributing to the thermoregulation of the animal. The new
paper demonstrated through colored scanning electron micrography how the internal walls of the hollow guard hair could not
conduct heat. This new knowledge of fur morphology does not
compromise the architecture developed from the original idea
because its functional performance as designed is still valid. The
active building envelope is covered with tube-like structures that
use a mechanized tracking system to follow the movement of the

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sun, maximizing heat absorption by the thick insulated envelope.
This mechanism is still valid, even if the science that inspired it is
no longer found to be true.

Namib desert beetle, Physasterna
cribiripes. (Photograph courtesy of
A. Sosio)

Another case study involves Namib desert beetles and their water
harvesting strategies. One particular species was studied for its
ability to collect water through a behavior known as fog basking. The study concluded that bumps on the dorsal surface of the
beetle have physical properties that allow water droplets to be collected and roll into the beetle’s mouth. It was later discovered that
the species the study focused on was incorrectly identified, and
it was not even based on a species that fog basks. The discovery
happened during the development of this book and led to the project being iteratively revisited, adjusted and rearticulated based on
this new scientific development. Each project’s inspiration follows
its own path, and what emerges from this is the importance of
the continued communication between science and design. We
attempt to teach an investigative interdisciplinary methodology
intended to provide new precedents for study while permitting
creative minds to make the leaps and lateral moves, necessary to
the design process.
The ability to understand the interrelation of systems is important
to all design professionals and can be enhanced by interdisciplinary education. Seeing everything as connected is an ability not
common to all, but something that can be acquired. Natural sciences are a great place where a designer can learn about connectivity despite some scientists’ tendencies to focus on detail. It is
not the discipline but the individual’s mind-set that matters. This
process happens through use of an interdisciplinary team where
each specialized member keeps an open mind toward discovery
and translation. The ultimate goal of our studies in this book is
not only to apply the organism’s features to design, but to gain
meaningful inspiration from the way in which that application
enhances creativity in the design process. It is not only the specific functional lessons that an organism conveys but the broader
themes of interrelatedness between components of nature that can
be applied to the architecture. Our way of working emerges from
a conscious collaborative effort to observe and then find design
concepts from two worlds that initially seem to have little in
common – the built environment and nature.

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While Part I discusses the biological and architectural background, Part II describes our methodology, which is applied to
twelve case studies. In particular, Part II looks at how architecture is inspired by four key biological functions: communication,
thermoregulation, water balance, and protection. Through initial
biological analysis and the development of the projects, the protoarchitectural proposals aim to provide a pivotal point in the relation between the built and un-built environment, providing a shift
from the exploitation of nature to exploration and collaboration
with it.

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“Consciously emulating Nature’s genius means viewing and valuing the
natural world differently. In biomimicry, we look at Nature as model,
mentor, and measure.”

- J.M. Benyus

Our approach to biomimicry seeks to expand on the most widely
used applications of the discipline to design and architecture.
Rather than mimicking or recreating nature through design, we
draw primarily from the functional, or performative, aspects
of nature and integrate those with the environmental context in
which such aspects are found. The natural world informs design
through a synthetic understanding of the adaptations an organism
has to its environment and the relationship it has with other organisms in the ecosystem in which it is a part. Using this approach we
can bring functionality to designed elements and thereby “inform
the form.” The resulting project is not a direct translation from a
particular organism, but is inspired by study of the function and
its context within a natural system. Novel and unexpected forms
consequently emerge from this explorative process. Though the
primary driver of the design is function, form is certainly not forgotten as it constitutes a fundamental aspect of design. Humans
are instinctively drawn to the complex, diverse and elegant forms
from the natural world. Through these projects, by beginning the

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process with the performative aspects of nature, we attempt to
ultimately achieve beauty. In this way, nature is a great model.
Bio-inspired design can help achieve the goal of lessening our
impact on the environment. Rather than creating a distinction
between the built and un-built worlds, our approach interrelates
the two, recognizing that all systems in the natural world are
continuous and coexist with each other. Rather than inserting
architecture into nature, our goal is to shift this perception and
integrate built forms into the natural world. The merging of
biology and design leads to discovery, innovation, and a novel
position in our relationship to the environment. This paradigm
shift will only occur by initiating a dialogue in contemporary
architecture that moves beyond pure formal and sustainability
concerns and aspires to connect directly with nature on a performative level. Our proposed method offers one way of exploring
biomimicry through functional analysis, and allied disciplines
are taking a similar approach. Parametric design, for example,
has renewed interest in the complexity of nature and processes
governing natural phenomena. Advanced technologies look to
the natural world to help modify how products are manufactured
and provide comfort to human habitations. As society moves
toward an environmental consciousness, we are encouraging
the use and understanding of nature to reformulate our aesthetic
sense. To illustrate our methodology we have used examples
from the animal kingdom as a source of inspiration. We considered the climatic and ecological context in which animals
evolved, and we drew upon those adaptations that have allowed
them to become successful in those contexts. Specifically, we
focused on animal skins, translating the observed adaptations
into the built environment. Skin offers one of the major models
or sources of inspiration that architecture can draw from biology.
There is an abundance of other models, and the book’s method
should be used as a way to guide oneself through the process in
an analogous fashion. We use the term “skin” on a general level
to refer to any animal covering, including fur, feathers, scales,
exoskeletons, and shells. Animals possess remarkable variation in the type of skin they have and in the ways they use the
critical functions of skin, such as protection, sensation, and heat
and water regulation. To draw an architectural parallel, building envelopes serve multiple roles, as interfaces between building inhabitants and environmental elements (e.g., water, air, sound,
light and temperature). That is why the word “skin” is often used to

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refer to building envelopes. However, while recognizing the parallels between natural skin and building envelopes, we strive to
avoid simplistic interpretations, such as attributing anatomical
characteristics to buildings.
The work presented here is a series of proto-architectural projects that focus on the development of the architectural envelope.
These projects explore the essential meaning of a building and
its envelope as they investigate the elements that provide shelter, protecting inhabitants from external forces while creating an
interface with the environment. The resulting projects perform
and respond; they take into consideration the dynamic local environmental conditions, enhancing and supporting these conditions
rather than exploiting them. Consequently, the focus on building
envelopes greatly facilitates the creation of a more sustainable
way of building and living.
While animal skins constitute the main driver for the design
inspiration, the studies show recognition of the importance of a
systems integrated approach. Once the skin’s major performative
function was explored, we expanded our research to a physiological and behavioral analysis of adaptations that help an organism
succeed and survive. This examination led to other elements
important to the architectural resolution of the design. The process
starts with the fundamental boundary element, the skin, with the
understanding that in order to design a coherent envelope, which
in nature and architecture comprises a system, it is important to
describe the elements, connections and interactions between the
components of that system. Animal skins and building envelopes
are a departing point for our approach to bio-inspired design. As
boundaries, they provide an essential element for connection and
integration of inside and outside, of built and unbuilt.
The following chapters are organized around four important skin
functions: communication, thermoregulation, water balance and
protection. Each chapter has two parts. The first introduces and
analyzes the selected function, illustrated using animal examples.
The second presents a few proto-architectural projects based
on in-depth analyses of individual animal skins and associated

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The book draws examples from the animal world that exhibit the
four major functions of skin we have chosen to examine. Our thesis is that through an evaluation of how the environment influences the anatomical, behavioral and physiological adaptations
of animals, we can develop a deeper knowledge of how human
constructions are affected by, and affect, the environment. This
process results in an increased design intelligence derived from
studying a range of investigative opportunities selected from the
observation of nature. Only limited components of both the animal and the design are shown, allowing for an openness of interpretation and exploration of the remaining aspects.
Rigorous and systematic analyses of the performative aspects of
the animal skin are the initial steps of the proposed methodology. Climate and habitat data are taken from the scientific literature, and the organism is analyzed in its environmental context.
Subsequently, drawings based on observations of the animal are
made in order to formulate possible architectural applications.
The analysis of the animal is conducted through line drawings,
diagrams, and renderings of the organisms to yield a synthetic
yet distilled portrait of the architecturally relevant adaptations.
The architectural rendition of biology is simplified and selective
as it draws from biological solutions to environmental challenges
in nature and describes them in a language that traditionally pertains to architecture and design.
As the proto-architectural project is developed, the scientific process is combined with the creative process. Though the scientific
process requires creativity, it tends to function in a more stepwise and iterative manner, while the creative process is characterized by leaps and returns; it is labyrinth-like and can appear, in
fact, quite “messy.” We allow for and encourage this nonlinearity, while simultaneously ensuring that the design is still fundamentally based on a platform of biological information. During
the proto-architectural exploration, the bio-inspired research is
consistently referred to, and the biological concept is present in
the ultimate design. The proposed methodology enhances and
strengthens the links between the biological and the architectural

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The challenge likely to be encountered with this truly interdisciplinary methodology is the necessity to build a common linguistic platform for communication. It is important to recognize the
need for the time it takes to develop a shared vocabulary and tools
to lead to effective comprehension and information exchange and
reduce the semantic noises common to such work. This process
will be different for each team and each task, and it is likely that
a common language needs to be developed for each goal. In our
case the challenge has been greatly facilitated through drawings.
They help identify, evaluate and define the knowledge relevant to
the overall objectives. The generation of a common visual language has allowed for enhanced opportunities for innovative solutions to be created between disciplines.
Both the biological elements and architectural ideas are explored
using the most diverse visual tools available. First, line drawings
are created to isolate key elements by indicating the functions that
are being examined. Taking the animal’s body as a starting point,
the drawings grow step by step like architectural plans and sections, retaining the scale of the original and concentrating on one
part of the body to define it in detail. The aim is thus to create a
stratigraphic sectional exploration of the skin, the primary organ
of investigation and design inspiration. The diversity of tools
ranges from hand-sketched drawings to 3-D digital modeling,
physical models, and digitally fabricated mock-ups. A process of
initial research leads to a codified design through a process of
graphical analysis and design synthesis. The final rendition of the
designs are proto-architectural projects rendered to substantiate
the final conceptual stage.
The projects explore the insertion of the design element in a particular setting so as to arrive at the material detailed definition of
a wall section, which aspires to describe the material components
and their performative characteristics. The basis for the design
might contemplate one specific animal trait or might emerge
through the synthesis of multiple traits. At times the propositions
that develop may not directly coincide with the initial feature analyzed; however, they potentially engage and result from its form or
function applied to other aspects of the design. A building envelope is often constituted of multiple parts that are either layered or
otherwise assembled to provide the functional qualities required.
These parts are interrelated, similarly to the way in which the
skin interrelates with other systems in living organisms; the way

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they are connected implies a multiplicity of functions that are in
The following twelve projects exemplify the conceptual framework outlined here. Each design project is organized into ten
parts (pages) to explain the biological research and inspiration
and subsequent development of the architectural project.
Project Parts
Animal Examples   The animals introduced on this page share
similar characteristics with the focal project species. The examples listed are then used to further demonstrate one of the four
major skin functions.
Taxonomy   Each project focuses on one species due to the qualities it exemplifies in relation to the chosen skin function.
Habitat & Climate   The habitat, climate and ecological context of the animal species are described. Selected variables are
detailed, including temperature, precipitation, humidity and wind
patterns. This information is particularly relevant as an introductory element to the biological analysis as well as the architectural
Animal Physiological, Behavioral & Anatomical Elements 
Major adaptations applicable to the four skin functions under
study are introduced with structural, physiological and behavioral
traits highlighted.
Interface between the Skin & External World  The type of skin
covering is described as well as the components that comprise
the skin. Aspects of the skin that exemplify the functions being
studied are shown.
Interrelationship between the Skin & Internal Systems  A
description of the internal physiological systems that interface
with the skin is given. The systems were chosen to complement
the major skin attribute of the species chosen.
Proto-Architectural Project  The design program and location are introduced. The main design drivers and strategies are

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shown through diagrams that describe environmental forces and
scenarios (for example, day/night, summer/winter, and rainy/dry
seasons) critical to the understanding of the envelope’s development. The location chosen is within the habitat of the animal.
Assignments were given in a didactic manner to facilitate the
design process.
Project Documentation  Diagrams and details derived from the
biomimetic aspects of the design are rendered, such as modular,
tessellating, and aggregating strategies.
Section of the Building Envelope  A wall section shows the
articulation of the key project components and describes the
entire wall assembly in its performative and material aspects.
Project 3-D Rendering  The final rendering offers an atmospheric and phenomenological overview of the project.
Skin Composition and Functions  The diversity of animal
skins provides fertile ground for bio-inspired design. Animals
have adapted to a wide range of environments, including those
extreme in temperature and rainfall, and animal groups have
unique strategies for dealing with the climate they exist in.
The skin is an ideal organ to use as inspiration in architecture
because of its multifaceted functions. It performs multiple, complex tasks yet it is one definable and readily visible system of the
body. Skin is also a duality. It constitutes the threshold between
the interior and exterior realms; it is the element of connection
between the two. Skin is a barrier, yet it is permeable. These comparisons are true and apply to both the biological and the architectural worlds. Using skin with its many parts and functions as
inspiration for the design of building envelopes allows us take a
holistic and systems-based approach to the proposed projects.
The skin performs numerous critical functions. It acts as a protective barrier against pathogens, predators and the environment.
It aids in temperature regulation through mediation of heat gain
and loss. Skin regulates water balance in the body through preventing its loss and by storing it or releasing it. It is a permeable
barrier that allows essential elements like oxygen and nitrogen
to diffuse into it. Nerve endings that respond to temperature and

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pressure are contained in the skin, leading to changes in sensation. The patterning and coloration of skin serves important communication functions. Through all of these diverse functions, the
skin can serve as a starting point for innovation, expansion and
enhancement in architectural design.
Skin is technically the soft outer covering of animals. However,
in this book the term is used in a much broader sense to mean any
animal covering. Examples of organisms with fur, feathers and
scales are used, because these are structures that originate from
the skin. Any structure that encloses an animal’s body and serves
protective and regulatory functions provides a point of departure
for our explorations.

Close-up of human skin. (Photograph
courtesy of S. Yelisee)

To introduce details of the structure, function and physiology of
skin we have chosen to describe human skin, because it provides
an easily understood model.10 Human skin is composed of three
major layers: the epidermis, the dermis, and the hypodermis. The
epidermis is a thin surface layer that acts primarily as protection.
It contains keratin, a protein, which keeps water in and harmful
chemicals and pathogens out. Specialized cells called Langerhans
cells engulf invading microorganisms and send messages to the
immune system for activation. Melanin, a pigment, is produced in
the epidermis. This pigment proliferates in reaction to UV light
and protects deeper tissue from sun damage (the process of tanning). The epidermis also functions in preliminary vitamin D
production, where chemicals produced interact with UV light,
leading to remote formation of the molecule that promotes calcium absorption. Too little vitamin D can weaken bones, and thus
this process is critical for human health.
The layer beneath the epidermis is called the dermis. It functions
in blood circulation, thermoregulation, protection from stress and
strain, and sensation. The dermis allows blood vessels located in
the lower skin layer to bring oxygen, water and nutrients to the
epidermis, where growing cells are fed through diffusion and
osmosis. When the body is hot, blood vessels and capillaries dilate
and conduct heat to the epidermis for surface cooling through
radiation and sweat production. When cold, vessels constrict and
retain body heat. In addition, nerve endings and special receptors
are located in the dermis that allow the body to respond to heat
and touch sensations. A thick layer of tissue within the dermis has
collagen and elastic fibers, providing insulation, cushioning and

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A generalized depiction of the layers of human skin. (Image courtesy of im
studio mi/la)

elasticity. Hair follicles and sweat glands are also located in this
Below the dermis the hypodermis is found. It is composed of a
thick layer of loose connective tissue containing larger numbers
of adipose, or fat, cells, which store energy. Major blood vessels
also occur in this layer. The hypodermal layer provides insulation,
acts as a shock absorber, and allows the skin to slide smoothly
over muscles, bones and joints.
As well as integrating with the major internal body systems, such
as the circulatory and nervous systems the skin also interrelates
with the digestive system. The digestive system is crucial for skin
function as the intake and digestion of fats and essential oils facilitate and maintain the skin’s function as a protective barrier, as
well as nourish glands and hair follicles. Vitamin D production
by the skin helps the uptake of calcium from food introduced into
the alimentary (or gastrointestinal) system. In the nervous system
the skin acts as the initial source of input, the signals of which
are transferred to the brain, for senses such as pain, touch, temperature, pressure and vibration. Nerves in the skin also instruct
skeletal muscles to shiver to produce body heat.

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(Top) Feline fur.
(Middle) Amphibian scales.
(Bottom) Pennaceous bird feathers.
(Photographs courtesy of
I. Mazzoleni, D. McShaffrey,
L. Mazariegos)

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Vertebrate groups such as mammals, amphibians, reptiles, fish,
and birds have multiple layers of skin that function in ways similar to human skin even though it may not be immediately obvious. Many of these groups have various protuberances originating
from the skin, called appendages, which work in conjunction with
the skin. While humans have small amounts of hair on their bodies, most other mammals have fur that serves to thermoregulate,
protect, sense, and communicate. Like human hair, fur is composed of keratin although it is usually made up of two layers —
ground hairs and guard hairs. Ground hairs are the bottom layer
of fur that tend to be thick and serve as insulation. Guard hairs, the
top layer of fur, are usually longer and coarser than ground hairs.
Guard hairs contain pigmentation, which can help in camouflage
or in attracting mates through coloration patterns. This layer also
serves as protection from elements such as rain, because it often
has water-repellent properties. Fur further aids in thermoregulation through nerves in the skin that respond to cold and heat.
The nerves activate muscles in the hair follicles that contract to
pull the hair shafts erect, creating insulating air spaces. When the
shafts are flattened, less air is trapped in the fur and the animal
is able to release heat. In some cases animal fur is modified into
hard, spiny protrusions that can serve as further protection from
predators. Fur also delivers sensory messages to the body. Other
appendages originating from the skin include scales and feathers,
which are also make of keratin. Scales are hard plates that serve
in protection from the elements, predators, and even prey. Snakes
and lizards are the animals most associated with scales; however,
a particular group of mammals — pangolins — has scales as
well, though they are actually modified hairs. Feathers, on the
other hand, are an appendage unique to birds; they are often considered the most complex appendage in vertebrates. There are two
main types of feather that serve somewhat analogous functions as
the fur of mammals. Pennaceous feathers have hooks and barbs
that lock together to provide a solid, stiffened surface for flight.
Birds coat their feathers with wax secreted from a specialized
gland that functions in waterproofing, through the conditioning
of feathers, and parasite resistance. Plumaceous feathers do not
have hooks and barbs and are therefore fluffy, allowing air to be
trapped and provide insulation.
The term “skin” is not commonly used for the outer coverings of
invertebrates. For example, insects, arachnids, and crustaceans
all have what is called an exoskeleton. This is a hard structure that

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supports, covers and protects an animal’s body. It can consist of
a variety of compounds, but most often it is made of chitin. It is
somewhat comparable to the function, but not structure, of keratin. Due to their rigidity, exoskeletons have a number of functions,
including protection, sensation, a barrier to water loss, communication (through their coloration), and support. There is further
variation in skin types that will be further explored in this book.

Dung beetle (Scarabidae) exoskeleton detail. (Photograph courtesy of
S. McCann)

The Four Selected Functions  To fully explore the relationship between animal skins and building envelopes, the book
focuses on disparate animals, skin types, functions, and climates and attempts to demonstrate the variety of ways the
developed methodology can be implemented and expanded by
others. We have selected animals that show clear and, in many
examples, unusual adaptations to four major functions of skin:
communication, thermoregulation, water balance, and protection. These particular themes were selected because they are
the most interesting and relevant to application in architecture
and design. Thinking about how these functions are accomplished in nature can lead to innovative ways of providing
human comfort, while lessening the built world’s environmental impact through changing the ways in which we design.
Communication is crucial for animal survival, and strategies
for communication take many forms. Modes of communication provide great insights to architectural investigations as
designers learn from animals about the exchange of information within the built environment. We focus here on one form
of communication widely used in nature: coloration. Coloration
is used for warning, protection, camouflage, and sexual attraction. In architecture, some designers have explored the use
of color as a communication tool, but more can be done by
improving and diversifying the use of color through implementing the strategies of the selected organisms.
Thermoregulation is the ability most animals depend on to
keep their body temperature within certain critical boundaries. Extremes in temperature pose tremendous physiological
challenges to living organisms, and various mechanisms to
help regulate temperature have evolved. Endotherms internally maintain their body temperature, whereas in ectotherms,
temperature regulation is a function of their external environment. In either case, thermoregulation is achieved through

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remarkable physiological and behavioral processes. One of
the major challenges architecture faces is to provide thermal
comfort to human inhabitants, particularly in extreme circumstances, when human bodies cannot acclimate to external conditions. Animal systems have much to teach us regarding how
to control temperature as well as how to limit energy expenditure in doing so.
Water balance is crucial for all organisms, considering that cells
are composed primarily of water. Water is needed for numerous
biochemical reactions, and it can dissolve and transport nutrients
and other molecules. Animals have evolved many novel strategies
to collect water and prevent water loss, particularly in water-limited habitats. In design, learning how animals prevent water loss
may extend humans’ ability to survive in dry and inhospitable
conditions through implementing systems that minimize the use
of water as well as collect, store and reuse it.
Protection from predators, parasites, physical injury, and the
environment can occur in many ways. Humans can learn from
the novel and complex adaptations that have evolved in the animal kingdom. Shelter, the most archetypical element in architecture, serves to protect or shield inhabitants from many things, and
designers, by drawing inspiration from the efficiency of animal
systems and the multiplicity of functions integrated within one
system, can develop novel, responsive solutions.

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^ polar bear
^ snow leopard
banana slug ^ ^ side-blotched lizard
ochre sea star ^
^ sea slug
^ dyeing dart frog
^ violet-tailed

^ tree pangolin

Namib Desert ^

^ urania moth
^ hippopotamus

The animal species and localities chosen for the projects are shown on a
world map. (Image courtesy of im studio mi/la)

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Animal communication occurs when a signal sent by one individual has an effect on the behavior of at least one other individual.
Communication happens via all senses and can occur through
smell, touch, movement, gestures, sound, vibration, facial expressions, electrical signals, and coloration. These various forms of
communication serve many functions: individual or species recognition, courtship rituals, warning off predators, aggression and
territoriality, and signaling about food or other resources.

The velvet-purple Coronet hummingbird has dazzling iridescent plumage
used in courtship rituals and displays
of territoriality. (Photograph courtesy
of J. Rothmeyer)

Animal coloration patterns can be powerful signals, because they
aid in all aspects of communication. Colors are wavelengths of
light; differences in wavelengths produce different colors. Bright
and contrasting color patterns send the signal to predators that
animals are poisonous or distasteful and therefore harmful to eat.
Camouflage helps animals blend into their surroundings; prey can
use camouflage to avoid predators, while predators use it to sneak
up on prey. Certain coloration serves to confuse or dazzle predators. Many animals mimic the color patterns of other animals,
usually to avoid being eaten. Colors also serve to promote sexual
attraction and therefore reproduction.11
Animal coloration is due primarily to pigmentation. Pigments are
chemicals produced inside tissues such as the skin, skin appendages, and even internal tissues and organs. There are many kinds
of animal pigments; the most important in the color of animal
coverings are melanins, carotenoids, and pterins.12 Some specialized cells do not contain pigment but still function in coloration.13

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Coloration can also be due to microscopic changes in the physical
structure of an animal’s surface, or a combination of pigmentation and structure. The physical characteristics of the surface can
cause the scattering, interference, and diffraction of light, leading
to iridescence.14
Animals perceive color in many ways, and there is a great range
of ability to detect different colors (wavelengths) among animals.
Some animals can only detect the difference between light and
dark, while others have sophisticated eyes that perceive many
colors and sharp images, and have good depth perception. Many
aquatic animals have evolved excellent color perception, because
objects underwater are harder to distinguish based on contrast
against the background. Bees, some birds and perhaps a few
mammals can see UV, or polarized light, because objects in their
environment absorb it. Most birds see in color and, in fact, have
colored oil droplets in their eyes that help them have even better
color vision than humans. Most mammals, however, have limited
color vision; humans and other primates are the only mammals
with a well-developed ability to perceive color.

(Top) The bright colors and iridescence of a hummingbird’s feathers can serve to attract mates, to
display aggression, or to warn off
predators. (Photograph courtesy of
L. Mazariegos)
(Bottom) Males and females of the
cobalt blue tarantula have iridescent blue legs until they reach the
final stage of sexual maturity, when,
oddly, males lose their iridescence
and become brownish in color.
(Photograph courtesy of Flamesbane

Designers can find inspiration from understanding the relationship of an organism to its ecosystem as well as how humans perceive their color and how that perception can be useful in design.
Most organisms do not see color as humans do, so communication between animals cannot be duplicated by humans. However,
we are ultimately designing for the human perspective; that is the
priority of the design process. The case studies presented embrace
different strategies for coloration and are used for different architectural purposes.
Buildings communicate in a variety of ways, at times as status
symbols and visible icons and at other times blending in invisibly and passing unobserved. There are four aspects of particular
importance in design, two of which stem from modernism —
extreme building height and façade transparency — and two of
which are more contemporary and have been looked at in this
book — color and camouflage.
Historically, façades have been important elements symbolizing
power and strength, designed to communicate corporate identity and establish visual supremacy with their presence in the
city. Skyscrapers, the equivalent of Middle Age towers, are the

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modern icons of power. Starting with modernism the extensive
use of glass allowed these tall beacons to convey not only physical
transparency but also a conceptual one, a manifestation of corporate operational transparency for public shareholders. Exemplars
are Ludwig Mies van der Rohe’s New York Seagram Building
(1958) as well as the Chicago Sears Tower (1973) by Skidmore,
Owings & Merrill (SOM).
Beyond the use of transparency in recent decades, few architects
have ventured into using coloration as a means of communication.
Each project of Berlin’s Sauerbruch Hutton Architects uses color
as the primary means to establish an architectural presence in the
urban environment; the building massing is articulated with gradients of colors: greens turning into oranges turning into pinks,
leading the viewer through a “rainbow” that dissolves the building presence from a single mass into a fragmentation of individual
unique elements. To the contrary, Emilio Ambasz’s projects often
bury themselves under layers of earth, camouflaging and causing
the building to disappear under strata of grass, hiding away while
providing the benefit of thermal mass to the building’s inhabitants. In addition, these buildings often provide the city with a
green public space.

Panel facade composed of timber
and polychromatic glass. Sauerbruch
Hutton Architects, The Federal
Environment Agency, Dessau, 2005.
(Photographs courtesy of Sauerbruch
Hutton Architects,
and Annette Kisling)

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Nature can provide functional inspiration for novel uses of color
in design. By studying the reflective microstructure of the Urania
moth’s wing scales, a project was designed that uses discarded
colored plastic bottles as a signal to inform villagers when water
is available. The iridescence of the violet-tailed sylph hummingbird inspired a colorful pavilion made by interlocking highly
reflective panels to communicate about activities occurring in the
pavilion. The photosynthetic lettuce sea slug inspired a center that
uses different species of algae to produce biofuels while simultaneously providing a chromatically dynamic beacon.

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Animal Examples — Communication  The black and white
stripes of zebras can confuse predators that cannot see in color,
and also make them unappealing to biting insect parasites.
Leafhoppers and thorn insects use both coloration and appendages
to make them difficult to distinguish from their environments.
Some reptiles, such as the chameleon, are able to voluntarily regulate the size of pigment holding cells and to change color as they
move from place to place. The chameleon also uses this ability
to send signals to members of its own species. The bright coloration of the blue-banded goby appears to stand out to the human
eye, but in vividly colored reefs it is hard to spot. The leafy sea
dragon floats slowly through the water like a tangle of seaweed.
The predatory stonefish is colored and ornamented to camouflage
among the rocks on the sea floor as it lays in wait for passing prey.

Animals ranging from invertebrates to large mammals use coloration for a multitude of purposes. (Photographs courtesy of: zebra, J. Rothmeyer; blue-banded goby, M. Bartosek; chameleon, K. Tolley; stonefish, B. Larison; leafhopper,
S. McCann; leafy sea dragon, I. Mazzoleni; thorn insects, M. Hedin)

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Animal Examples — Communication  Flamingoes are naturally white, but develop a pink coloration as carotenoids from
their shrimp prey are incorporated into their feathers. Dragonflies
use color to warn off predators. The bright, colorful plumage of
the male mandarin duck is permanent and its intensity provides a
measure of his genetic condition to the dull brown females. Both
sexes of the scarlet macaw are boldly colored; this might facilitate
recognition among members of the same species in dark rainforests, but scientists are still unsure. The dominant male mandrill
in a group develops a brightly colored face that signals his status
and restricts breeding rights to him alone. The male anole fans
a brightly colored flap of skin called a dewlap from his throat to
advertize his presence and condition to females in the area. In
ladybugs, black spots stand out against a red background to warn
predators that they are distasteful.

Animals ranging from invertebrates to large mammals use coloration for a multitude of purposes. (Photographs
courtesy of: flamingo,; dragonfly, B. Larison; male anole & ladybug, S. McCann; scarlet macaw, A.
Kirschel; male mandril, Wikimedia; mandarin duck, M. Montese)

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Animal Examples  Many insects possess striking coloration patterns; the function of these is likely to ward off predators or attract
mates. The saddleback caterpillar is bright green with a white and
brown circle resembling a saddle. They have irritating, venemous
hairs on the protrusions on the front and back of their bodies, which
prevent them from being eaten by predators. Cuckoo wasps have
vibrantly colored metallic bodies, leading to the additional common names of jewel wasps or emerald wasps. Many beetles are
iridescent, including the family called the metallic wood boring
beetles. This family has often been used in human ornamentation
and decoration because of their dazzling coloration. Glasswing
butterflies have partially transparent wings. This may allow them
to camouflage from predators by blending in to their surroundings.
The physical mechanism that causes transparency is not known,
but may be due to having fewer scales.

Insects possess a lot of variation in coloration, including having bright, iridescent or even transparent coloration.
(Photographs courtesy of: saddleback caterpillar, cuckoo wasp and metallic wood boring beetles, S. McCann; glasswing butterfly, S. Yanoviak)

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Urania moth

Chrysiridia rhipheus




C. rhipheus

Photograph courtesy of A. Richards, Bohart Museum of Entomology, University of California at Davis.

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Habitat & Climate  The Madagascan sunset moth only occurs on
the island of Madagascar. The island’s climate varies considerably
longitudinally. The eastern part of the country supports a rainforest habitat due to trade and monsoon winds along the east coast
causing high precipitation levels. The winds lose moisture while
moving west, so the central part of the island is drier; it is also
cooler due to the higher altitudes. The western part of the island
is mostly arid, with semi-desert conditions in the southwest. Two
principal seasons are defined: a hot, rainy season occurring from
November through April and a cooler, drier season occurring from
May through October. The sunset moth is present throughout most
parts of Madagascar, but they migrate across the country throughout the year in response to changes of the host plants they depend
on for survival in the early stages of life.




Dry deciduous forest in the western region of Madagascar. Close up of Omphalea’s leaves, the moth’s host plant.
(Photographs courtesy of A.P. Raselimanana)

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Animal Physiological, Behavioral & Anatomical Elements 
The behavior of C. rhipheus is atypical of many moths, because
it is active during the day. The coloration patterns on the moth
are warning signals to let potential predators know the moths
are toxic. The toxicity comes from trees and shrubs in the genus
Omphalea. Caterpillars feed on the toxic plant components,
which are stored through their development, even in to the adult
The dependence on the host plant is so strong that sunset moths
migrate to different populations of their host plant during the
year; specifically, they migrate from species in the dry forest in
the west to the rainforest in the east. The moth caterpillars negatively affect the Omphalea populations, because they eat the flowers and fruit from the entire plant.










C. ripheus caterpillars feed exclusively on Omphalea plants, often completely defoliating them. They eat the leaves,
flowers, fruit, tendrils and new shoots, causing damage to all parts of the host plants. (Photograph, Wikimedia; project team: B. Frati & N. Zarfaty)

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Interface between the Skin & External World  The most
striking feature of the Madagascan sunset moth is its multicolored wing patterning, including areas of brilliant iridescence.
Iridescence is defined as the surface of a structure that changes
color when the viewing angle or angle of lighting is changed.
Iridescent colors are structural colors; they tend to be brighter
and purer than colors produced by pigments. Butterfly and moth
wings are comprised of thousands of partly overlapping scales,
and the structure of the scales affects the way they reflect light.
Sunset moths have two layers of scales on their wings: the inner
layer and cover scales. Each cover scale is a multilayered structure made up of thin layers of cuticle between thin layers of air.
The layers of cuticle are supported by very small, spaced out
columns of cuticle. The layers of cuticle and air can be different
thicknesses, which create the variation in color in the wings.



Scale’s Curvature



Scale Multilayer Microstructure

In iridescence, which is caused by structural modifications, the hue of the color changes with the angle of the
observer and light shining on the surface. The brilliant colors are caused by the reflection and interference of light
going through multiple surfaces. (Photograph courtesy of A.P. Raselimanana)

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Interrelationship between the Skin & Internal Systems 
Different pigments can occur within scales, and with the combination of the multilayered cuticle surface, different iridescent
colors can be obtained. When light shines through the first layer
of cuticle, some is reflected back from the surface and some goes
through to the next level. With each level the light penetrates,
more light is reflected from the next level down. The reflected
light from each level travels the same way, that is, away from the
cuticle layers.
Light occurs in wavelengths, and if the crests of the wavelengths
overlap with each other, they amplify the effect of reflection.
When the wavelengths line up with each other intense coloration
occurs. This phenomenon is called constructive interference, and
it is what causes iridescence.

Chitin layer
Curved scales
Flat black scales

Detail of one curved scale

Microstructure of the curved scale
(Air cuticle)

In the panels the white arrows represent the direction of light shining on the scales and illustrate how wing coloration
is a function of angle of perception. The wings of the moth are made of two layers of scales: an upper, curved layer
and a lower black layer of flat scales. Both layers of scales are attached to a chitin layer. The microstructure of each
curved scale is responsible for producing the brilliant iridescent colors in the sunset moth. (Micrographs courtesy of
Yoshioka and Kinoshita [2007])

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Proto-Architectural Project  The project is located in the
moth’s natural habitat, an arid town in western Madagascar. The
design for the building envelope was inspired by the reflective
and refractive properties of the moth’s multilayer and multi-color
wing assembly. The building provides a new system of clean
drinking water for the local community where rainwater is collected through a large roof surface. A filtration system which
forces water from the roof into the building’s envelope is constructed to support the aggregation of reusable colored bottles
that collect water through a structural trellis. The angle of the
bottles changes depending upon the amount of water collected
in the bottles: when empty the bottles are perpendicular to the
façade and as the container becomes heavy with water it shifts
parallel to the façade. Light plays with the shape and angle of the
water-filled bottles, creating spectacular moments of reflection.

The coloration effect provided by the plastic bottles was analyzed through a series of experiments investigating the
relationships between a surface, material, water, light and the resultant colors.

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Project Documentation  The façade communicates water availability by displaying different parts of the colored bottles. The
base of the bottle is black, while the remaining part is colorful,
either blue or green, typical colors used to enhance passive solar
purification processes.
The bottles’ layer is sequenced to take advantage of the play of
light through the bottles’ curvature to create varying moments
of reflection. This layering technique, together with the overall
circular geometry of the structure, alters the angle of reflection
as each bottle has a unique position in relationship to the sun,
thus providing a kaleidoscopic effect. The façade explores both
the idea of color as a useful means of communication and the
physical properties of light and color.

Color is used to attract people to the building and is created through the superimposition of different façade layers,
with the light passing through the changing angles of the water-filled multi-colored bottles.

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The curved surface of the tubular structure and the variable distance between tubes is responsible for reflecting and
refracting color, which varies through time to announce water availability. The project proposes to reuse locally
collected material such as plastic tubes, pipes and bottles, an environmentally conscious and low budget strategy.

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Water scarcity and pollution are dire problems in developing countries. Finding simple ways to collect and purify
rain water is crucial to the well-being of these communities. Additionally, re-purposing the uses of plastic bottles, by
extending their life cycle, minimizes landfill and illegal toxic burnings.

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Animal Examples  Communication is not always delivered
using aural mechanisms and many organisms communicate via
visual display. Birds in particular have mastered visual communication. Bright hues and iridescent structures can act as a
signal of quality, territory defense, or even as a distraction to
predators. The violet eared hummingbird uses purple and green
iridescence created by alternating layers of refracting light striking pigments to attract females. The resplendent quetzal flashes
bright blue, a color created by light reflecting off trapped water
molecules within the feather. The fiery throated hummingbird
combines these cooler colors with bright reds and oranges created by physical pigments in the feathers. The swallow-tailed bee
eater combines these feather colors with a striking red eye pigment to capture an observer’s attention, be it a fellow bee eater
or a human.

The variation of colors in bird feathers acts as a form of communication vital to attracting the attention of the opposite sex, as a warning to competitiors, or even as a camouflage in a dense forest. (Photographs courtesy of violeteared hummingbird, resplendent quetzal, fiery throated hummingbird and swallow-tailed bee eater, J. Rothmeyer)

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Violet-tailed sylph

Aglaiocerus coelestis


Chordata Family:
Aves Genus:
Apodiformes Species:

A. coelestis

Photograph courtesy of J. Rothmeyer

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Habitat & Climate  The violet-tailed sylph (Aglaiocercus coelestis) can be found along the western slopes of the Andes, but is most
abundant in the cloud forests of Ecuador. These types of forests
are moist, due to the high level of precipitation that falls yearly;
the rainy season from January to May is cool, 8–15ºC (46.4–59ºF),
due to the high altitude, from 500 to 4000 m (~1640–13,123 ft).
This combination of moisture, 50 to 1000 cm/year (19.5–393.5 in),
and elevation creates slow evaporation, resulting in a lush, densely
forested habitat. The eastern and western slopes of the Andes in
Ecuador and Colombia experience rainfall fed by moist air coming from the Amazon Basin. Precipitation is high due to winds
that bring in water vapor from the Amazon. Such a unique climate allows for a variety of exotic plants to grow and flourish. The
Heliconia is one such plant, which produces nectar that the violettailed sylph feeds on.


The violet-tailed sylph has a home range that includes forest edges, shrubs, and deep cloud forests within Ecuador
and Colombia. This species is unique in its altitudinal range and is most easily spotted around 1000 meters above
sea level. These cloud forests are unique within the Andes and harbor much biodiversity and endemic species.
(Photographs courtesy of J. Rothmeyer)

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Animal Physiological, Behavioral & Anatomical Elements  A
hummingbird’s normal body temperature runs around 40.5ºC
(105ºF). When a hummingbird sleeps, this temperature will drop
to as low as 21ºC (70ºF). Due to its small size, the hummingbird rapidly loses heat to the outside world because of the ratio
between its volume (small) as compared to its relative surface
area (large). This fact, combined with the high, cool elevations
where these hummingbirds often reside, makes heat conservation
a priority. To conserve energy a hummingbird will go through a
nightly rapid transition in metabolism and activity equivalent to
a brief hibernation. This state is called torpor and is so extreme
that one can pluck a hummingbird off a branch while in torpor
without the bird even moving! Always on the search for energy
sources, many hummingbirds are nectavores and seek out this
high-energy food source even in the most protected of flowers.



Downy feathers mixed among
contour feathers
Filoplume feathers mixed among
contour feathers
Rectrices (tail feathers)

The most brilliant iridescent colors are found in males and have two purposes: to advertise their physiological quality
to other males and to attract females. Feathers carry sub-microscopic structures that produce bright colors; however,
the brilliance can be seen only with certain positions involving the bird, sun, and observer. (Photograph courtesy of
J. Rothmeyer; project team: J.M. Helinurm & A. Amiri)

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Sun angle



Interface between the Skin & External World  Hummingbirds’
feathers are functionally similar to other birds, yet produce unique
iridescent colors. The shaft of the feather (or rachis) extends out
as alternating thread-like barbs and barbules, which grow out
at proper angles in order to hook with one another and maintain structural integrity, but at the same time remaining flexible
enough to bend or disconnect without breakage. Feathers lie on
top of each other, keeping only the iridescent tips visible. The
keratin protein found in feathers provides flexibility and integrity,
and acts as a natural insulator. Contour feathers cover the body
and appendages of birds. These feathers have an expanded vane
that provides the continuous surface necessary for flight. Changes
in the angles of these feathers allow for changes in the amount of
feather brilliance observed from different positions. Some feathers are flat; others are curved.




Some feathers, for instance on the gorget and crown, resemble flat mirrors, and light that hits them can be reflected
in only one direction. The sun, observer (bird or human), and bird must be aligned properly to view the brilliance of
the plumage. When no light shines on these feathers they appear black. Other feathers, like those on the back, are
curved inward to resemble concave mirrors. These curves come in many angles, allowing for light to be refracted
to the observer at different directions. The section shows feathers insert into bird skin through the epidermis and

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Interrelationship between the Skin & Internal Systems  To
be as lightweight as possible, most of the hummingbird’s bones
are extremely porous, and their legs are extremely small, short,
and stubby. Perhaps the most diagnostic character of the hummingbird is its wing structure, which enables it to hover, fly backwards, and to change direction with remarkable precision. Several
hummingbirds can even fly upside-down for a short period of
time. The shoulder joint is a ball and socket joint that allows the
hummingbird to rotate their wings 180º in all directions. When
hummingbirds take flight, they move their wings in an oval pattern and maintain their body in an upright fusiform position, with
their entire body facing the world. When they are hovering they
move their wings in a figure-eight motion. A hummingbird can
fly at an average speed of 40 to 48 km/h (~25 to 30 mph) and dive
at a speed of up to 96.5 km/h (60 mph).

Upward flight

Forward flight





Backward flight

The black tip of the
feather indicates the
area where
iridescence is found.
Near the skin of the
hummingbird, the
shaft of the feather is
uncolored. When the
feathers lay on top of
each other, the area
that is uncolored
remains hidden.


Hummingbird wings beat about 70 times per second while in regular flight, and up to 200 times per second when
diving. To power this movement, their metabolism is extremely fast, and heart rates average 250 beats a minute, with
a maximum heart rate recorded of 1260 beats a minute. (Photograph courtesy of L. Mazariegos)

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Proto-Architectural Project  The pavilion is located in Parque
la Panecillio in Quito, Ecuador, a city 2800 m (9185 ft) above sea
level and 24 km (15 mi) south of the equator. Taking advantage of
the subtropical, yet cool climate (due to its elevation), the pavilion
develops a porous envelope, enabling the elements to permeate
through. Inspired by the iridescence of the feathers, the pavilion takes advantage of the color palette found on the violet-tailed
sylph: blues, violets, and greens. Color operates as a device for
aesthetics and a significant means of communication. It serves
as an indicator for occurring and upcoming events by radiating
colors in response to dynamic atmospheric changes. Bright colors
are a device to express energy and enthusiasm, thus enhancing the
sensorial and spatial experience of the inhabitant – from within
and afar.

Iridescence is dependent on the position of the object, the observer and the light rays hitting the object. Inspired by
these factors, 3D digital modeling techniques were utilized to explore the articulation of individual panels in relation
to their rotational array – with the intent to enhance and maximize the reflectivity of sunlight, while minimizing the
quantity of artificial light sources at night. (Photograph courtesy of L. Mazariegos)

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Project Documentation  The pavilion’s panelization and joint
system is inspired by the hooked barbules in a hummingbird’s
feather and the implementation of principles of light refraction
magnified by water. One level of refraction is developed through
the panels’ materiality, composed of two layers of semitransparent resin and a thin layer of glass to provide additional qualities.
A second level of refraction is established by understanding how
light reacts differently when it hits and travels through water
– resulting in variations in the colors of light dispersed. The
hook joint creates small water pockets between the panels, thus
enhancing the desired light deflection. Resource optimization and
efficiency are crucial to the pavilion and its use as public space,
hence the exploration and strategic implementation of a repeated
module to produce communicative effects.



Light refracts exiting drop

Some light
passes through
Collected water

Light refracts entering drop



Colored light refraction

Mimicking the complex structure of feathers, the modules overlap, thereby magnifying the light refraction between
layers, each made of glass and colored resins.

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Hooking S-joint
Water pocket

The structure displays a broad spectrum of color variation throughout the pavilion. The modular panels are arrayed
to reflect and refract the sunlight at varying angles. The color brilliance is ultimately enhanced through the combined
effects of the semitransparent panels and water pockets. The panel joints form an S-like shape to interlock with one
another, forming a small cavity to trap water.

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The pavilion acts as a beacon during activities and events, becoming a glimmer of coloration when observed from
the city. The colorful variations of refracted light infiltrate the internal spaces, amplifying the sensorial experience
promoted by the physical environment.

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Animal Examples  Marine invertebrates use coloration for different purposes. Giant clams are vibrantly colored, often with
iridescent patches on their mantles, or body walls. They achieve
this coloration through symbiotic algae harbored in their tissues.
The algae are given a protected place to live, and in turn, the clam
ingests the sugars and proteins produced by the colorful algae. A
group of sea slugs, called nudibranchs, are known for their vivid
Color patterns, often contrasting, are thought to serve as warning
coloration to notify predators of their distastefulness or toxicity.
The blue-ringed octopus is extremely venomous. They are small
and normally camouflage with their surroundings. However, when
agitated, their skin turns bright yellow, and the blue and black
rings covering their bodies darken substantially when threatened.

Vivid colors are thought to serve as warning coloration to notify predators of their distastefulness or toxicity. Color
patterns are often contrasting, like red paired with white or black paired with yellow, to be particularly conspicuous
to predators. On the other hand, coloration can serve as a camouflaging mechanism through which certain animals
can establish symbiotic relationships with others. (Photographs courtesy of: giant clam, red sea slug (Chromodoris
sp.) and blue sea slug (Phyllidia sp.), B. Larison; blue-ringed octopus, J. Himes)

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Lettuce sea slug

Elysia crispata


Mollusca Family:
Gastropoda Genus:
Heterobranchia Species:

E. crispata

Photograph courtesy of P. Krug

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Habitat & Climate  The sea slug Elysia crispata, often referred
to by aquarists as the lettuce slug, can be found on island coasts
throughout the Caribbean Sea. Its habitat is limited to the shallow waters of coral reefs and mangrove lagoons, approximately
1.5 to 12 m (5 to 40 ft) in depth. They live in areas where water
currents are relatively weak and temperatures range from
20–30ºC (68 to 86ºF) . Water temperatures closely mimic air
temperatures in the Caribbean, with little change across seasons
during a given year; that is typical for tropical regions due to their
close proximity to the equator. Air temperatures are warm (20–
32ºC, 68–86ºF,) and relative humidity is typically high (50–90%).
Slight seasonal shifts do lead to rainy seasons in the tropics. In
Haiti, the specific island chosen for this study, this increased rainfall occurs primarily from May to July.


Elysia crispata lives on coral reefs and in mangrove lagoons of the Caribbean Sea, where it eats green algae and can
store the algal chloroplasts in its own tissues. It is often found “basking” in the sun in order to provide energy for
photosynthesis, making it solar powered. (Photographs courtesy of R. Ellingson)

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Animal Physiological, Behavioral & Anatomical Elements  A
unique characteristic of Elysia crispata is its ability to take functional chloroplasts (plastids) from its green algal food source and
store them, a process called kleptoplasty. The slugs can be found
“basking” in clear and shallow waters, absorbing the sun’s energy
to drive photosynthesis (the production of energy from light, water
and carbon dioxide). This allows E. crispata and some of its relatives to go weeks or even months without food, making them the
only known solar-powered animals. The common name “lettuce
slug” comes from the leafy appearance of its parapodia, two large
flaps that run the length of its body. Basking refers to the opening of these parapodia to absorb light, made more efficient by the
increased surface area of their ruffled, leafy form. Stored plastids
cause E. crispata to usually appear green, but vibrant shades of
blue, yellow and hints of red can often be seen as well.



Unfurling increases
sunlight exposure

E. crispata has a translucent body with a variation of colors. The rhinophores protruding from the back can range
from blue to red to green. The slug’s color is known to fade or intensify with the sequestration of chloroplasts within
its tissues. The more recently a slug has eaten, the more colorful it will appear. E. crispata has the ability to expand
its parapodia in order to capture more energy from the sun, or fold them in close to decrease its vulnerability.
(Photograph courtesy of P. Krug; project team: A. Munoz & R. Hopkins)

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Interface between the Skin & External World  The skin of
Elysia crispata is translucent. Visible colors therefore come from
internal tissues and organs or any material that might be incorporated into the skin. Translucent skin is particularly important
for kleptoplasty since radiant energy from sunlight must reach
internally stored chloroplasts. The leafy appearance of parapodia
is crucial for increasing the surface area of the skin, allowing for
efficient uptake of light for photosynthesis. The process of kleptoplasty causes the translucent skin to change in color from clear
or pale to a rich green (or even blue), much like the leaf of a plant.
Sea slugs have no shells to provide physical protection from predators. Therefore they rely on cryptic coloration, often blending in
to the green color of their algae food, as well as large amounts of
mucus produced by the skin. This mucus is distasteful to most
predators and can often be toxic as well.


Algae cell



The chloroplast, contained in the algae cells, is the organelle responsible for photosynthesis. The slug has a slimy
skin that provides no physical protection. Instead, it produces mucus that deters predators because of its bad taste
and toxicity. Because they often adopt the green color of their algal food, camouflage may also be a useful tool to
avoid predation.

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Interrelationship between the Skin & Internal Systems  Factors
that determine the color of Elysia crispata are tightly linked
to the system it uses to harness energy. Since functional chloroplasts are sequestered in its skin, E. crispata usually appear
green. Kleptoplasty causes the slugs to be green in much the same
way that chloroplasts turn the leaves of plants green. Much of the
light at wavelengths near the red and blue ends of the visible light
spectrum are absorbed and used for photosynthesis, while most of
the green light is reflected back to be observed from the outside.
The digestive tract of E. crispata runs throughout the body and
its vein-like patterning is often apparent, especially after a meal.
Chloroplasts are stored in close proximity to the digestive tract to
aid in the absorption of carbohydrates produced by photosynthesis. If E. crispata goes an extended period of time without food,
few chloroplasts are present and its color becomes very pale.






Above: Close up view of E. crispata showing individual chloroplasts (tiny green spots) sequestered in its tissues.
Below: Illustration of parts of the lettuce slug’s internal anatomy. (Photograph courtesy of P. Krug; diagram redrawn
from R. Ellington)

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Proto-Architectural Project  Based on the eastern coast of
Haiti, the building intends to fill the need for a dependable energy
source during the time of reconstruction after the 2010 earthquake disaster. The envelope system, inspired by the processes
that generate color in E. crispata, is a photo-bioreactor made of
layered plastic modules and tubes filled with algae.
Using locally abundant resources — ocean water, sun, and
micro-organisms — the building produces biofuel, enabling it to
change colors, ranging from red to blue to green, as the algae
matures. The growth and harvest of the algae provide a dynamic
beautification of the structure that communicates its level of
energy production.

The building’s coloration is obtained with the use of living organisms such as algae. By communicating the harnessing of energy through color change the enclosure produces a presence along the coast of the island.

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Project Documentation  Different algal species grow under
unique conditions; therefore, a diversification of the species
used allows optimization of biofuel production. As a by-product,
variation in color display is also achieved. A system of interlocking tubes brings water to the plastic modules, which are layered
depending on particular sun exposure requirements. At any given
time there can be as many as six different species of algae maturing to create biodiesel, each exhibiting a different color. The
weather and seasons play an integral part, as the sun dictates how
fast or slow the algal growth process occurs.
Additionally, tubes filled with one more bioluminescent species
of algae are intertwined to the structural elements, thus providing
a beacon at night.

Day 1: Sea water and Bacillariophyta algae
Morning: Dinoflagellate algal blooms
within structural tubes (for bioluminescence only).
Separate system than the inner bio-fuel module
Day 5: Young bacillariophyta
Dawn: Bioluminescence begins
to be perceived with diminishing light

Day 10: Mature bacillariophyta ready to
be processed into biofuel
Night: Bioluminescent algal bloom illuminates the structure
around each module, creating a fabric of glowing color. Movement
within these tubes allows for a constant bioluminescence.

Algae are living organisms that use sun, energy, and CO2 to photosynthesize. Mimicking the translucent affect of
the E. crispata skin, the building’s hollow tubular structure of the envelope contains bioluminescent algae, creating
a light source during the night. The diagram indicates a timeline of harvesting cycles. Images show day, sunset, and
nighttime building coloration scenarios.

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Water and algae


Bioluminescent algae




Harvesting and machinery

Networks of structural tubes support a series of interconnected modules which contain micro algae whose main
purpose is to create biofuel. The translucent pipes contain bioluminescent algae whose main purpose is to provide
bright coloration during the day and bioluminescence at night.

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The multiple layers of sacs enclosing micro algae and the tubular structure containing bioluminescent algae create
an array of ever changing coloration inside and outside the structure. The variation in color is produced both by
the different types of algae and by their different level of maturation, providing a dynamic and continuously varying
message of productivity.

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The large ears of the fennec fox are
endowed with a network of blood
vessels that can be used to flush
excess body heat. (Image courtesy
of D. Kim & J. Cambron)

07_Thermoregulation.indd 97

Maintaining an appropriate body temperature is critical for all
aspects of biochemical and physiological function, and animals
expend a great deal of energy controlling their body temperature.
There are two main strategies by which animals regulate their
body temperatures. Ectotherms, which include invertebrates,
fish, and reptiles, obtain most of their body heat directly from
the environment, and their metabolic rates fluctuate with ambient temperature. Endotherms, such as mammals and birds, produce their own heat through metabolic processes and are able to
maintain constant body temperatures. Almost all heat produced
is ultimately lost, making endothermy an energetically expensive
process. Consequently, endotherms need to eat high-energy foods
or feed very often.
Both endotherms and ectotherms use behavioral strategies to
manipulate their body temperature when the environmental
temperature is either too hot or too cold. Heat is lost and gained
through four major mechanisms: conduction, convection, radiation and evaporation. Conduction occurs when two objects of
different temperatures come into contact with each other. For
example, to gain heat an animal might lie on a rock that has been
warmed by the sun; to cool down an animal might wallow in cool
mud. Convection is similar but occurs when heat is transferred
through air or water. This often occurs when an animal is hot
and its body heat warms the air around it. As air passes over the
skin, the mass of hot air is pushed away from the body and allows
a cooler mass of air to take its place. Animals can seek out cold

13/12/12 11:11 AM


air or water currents to cool down in this way, and conversely,
find warm air or water to warm their body temperature. When
heat transfer takes place without physical contact this is called
radiation. Objects give off heat in the form of electromagnetic
radiation when their temperature is warmer than that of the surrounding environment. This means that animals can lose heat
when their bodies are warmer than the ambient temperature, and
they can gain heat from the sun when they are cooler. Finding
shade to cool down or lying in the sun to warm up (basking) are
ways of behaviorally thermoregulating using radiation. The only
way animals can lose heat when the air temperature is warmer
than their body temperature is through evaporation. Evaporation
is a cooling mechanism that occurs when heat is released from
the conversion of liquid water to water vapor. Animals cool themselves using evaporation through sweating and panting.
Due to their large body size and small
surface to volume ratio, elephants
retain a lot of body heat. Therefore,
they flap their ears, bathe in water,
and stand in shade to cool down.
(Image courtesy of T. Barsegyan)

An animal’s size, or more specifically, the ratio of its surface area
to volume, is very important when considering heat gain and loss.
As geometrically similar shapes get bigger, their surface area relative to their volume decreases. Because the surface is where heat
is gained and lost to the environment, larger animals are therefore better at retaining body heat. This body mass to surface area
relationship is why bigger animals tend to occur in cold environments and smaller animals tend to occur in hot environments.
When large animals occur in hot environments, they tend to have
less fur to facilitate heat loss. In addition to varying body size,
evolution has acted to change the surface area of animals through
creating longer and bigger extremities in hot environments and
decreasing extremity length in cold environments. In hot climates
longer extremities, such as longer limbs and big ears, allow more
heat to be lost to the air, and shorter extremities can help prevent
heat loss, thus conserving heat.
Animals that live in extreme heat and aridity have evolved many
strategies to prevent overheating and to dissipate any excess body
heat. These range from the physical attributes of the animals to
physiological adaptations and behavioral changes. Many animals
in hot climates are light colored to reflect heat. Long limbs not
only allow for increased surface area but reduce radiation by keeping the body farther from the hot ground. Animals that cannot
sweat may rub saliva over their bodies or urinate on themselves
to promote heat loss via evaporation. Some animals have many
blood vessels near the surface of their skin to promote convective

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In cold environments ducks behaviorally thermoregulate by fluffing their
feathers, tucking one leg under their breast feathers, and placing their bills
in feathers. These strategies prevent heat loss. (Photograph courtesy of A.

heat loss. These vessels can dilate, or open further, to allow for
increased blood flow and more heat to be dissipated. Some animals allow their body temperatures to rise with the environmental temperature during the day and then allow the stored heat to
dissipate in the cool of the evening air. Many desert animals are
only active at night or at dawn and dusk and stay in protected,
shaded areas during the day. Small animals, particularly rodents,
lizards and insects, may take refuge in underground burrows to
prevent overheating in the hottest part of the day. Some rodents,
frogs and arthropods are able to save energy by going into a kind
of dormancy called estivation during the hottest months of the
In contrast to the hottest climates, many animals are adapted to
live in arctic environments where temperatures are extremely cold.
Most arctic animals have some type of thick insulation in the form
of fur, feathers, or blubber, a layer of subcutaneous fat. Fur and
feathers can be adjusted to trap pockets of warm air close to the
skin or release it when conditions are warm. Animals also curl up
when at rest to minimize heat loss through reduction of surface
area exposed to the environment. Under the skin, blubber prevents
heat from being lost from the body core to the environment. Some

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10 0

Lizards behaviorally thermoregulate by basking to acquire heat or retreating
into shade to prevent heat gain. (Photograph courtesy of S. Yeliseev)

birds and mammals have a complex counter-current heat exchange
system that allows warm blood coming from the heart to heat
cooler blood coming from the extremities. This system may also
occur in hot environments to cool blood flowing to temperaturesensitive body organs like the brain. Many ectotherms such as fish
and insects survive subzero temperatures by producing proteins
that prevent ice crystals from forming in their bodies. Additionally,
endotherms such as mammals and birds may dig dens or huddle in
groups to protect themselves from cold winds and temperatures.
Just as desert animals reduce their metabolic rate in estivation,
cold-adapted animals do the same thing when hibernating. This
strategy minimizes heat loss by decreasing the difference in temperature between the animal and the environment.
Building envelopes are responsible for the mediations between
indoor environmental comfort for the inhabitants and exterior
climate conditions. In order to minimize the use of natural
resources several passive strategies can be adopted. The most
important of all is the placement of a building on a site and its
orientation to environmental forces which can facilitate the use of
rain, sun, and wind to achieve internal comfort.
For most of human history - and in a few populations today nomadism characterized our way of life. This transient lifestyle
was based on the need to find food which necessitated periodic

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movement to adapt to the changing seasons and food sources.
Consequently, in nomadic societies shelters were not immobile;
they had to be portable or relatively easy to construct. These
kinds of structures, in comparison to the fixed ones most prevalent today, were less impactful on the environment. The rise of
agriculturally-based societies led to a way of life in which human
dwellings are designed for permanency, and thus the need to deal
with the year-round weather conditions of a location.
For example, in desert regions thick walls provide thermal mass
to help offset the high temperatures of the day while releasing
heat during the cool nights. Conversely, in hot and humid climates, such as the tropics, raised, light structures allow for maximum cross ventilation, optimizing evaporation.
The excessive use of active, energy intensive systems places an
undue burden on the planet. Passive strategies offer a valuable way
to reduce our ecological footprint. This will help us achieve energy
and carbon efficient buildings, more respectful of the environment and less demanding on the use of the planet’ finite resources.
Moreover, technology plays an important role in the implementation of environmentally friendly strategies. For example, sensors
can track and trigger appropriate adaptive responses in building
envelopes to changing external conditions. In addition, smart
materials capable of storing heat and even regenerating themselves
over time can be used for the optimization of natural resources.
The envelope made by ethylene
tetrafluoroethylene (ETFE) panels is activated using pneumatic
mechanisms triggered by weather
sensors. Cloud 9, Media-TIC, Barcelona, Spain, 2011. (Photograph
courtesy of A. Suner)

07_Thermoregulation.indd 101

The case studies presented embrace different strategies for thermoregulation in both hot and cold environments. The project
modeled on the side-blotched lizard combines behavioral and
physiological strategies to provide a comfortable living unit in the
fluctuating daily temperatures of the desert. The unit accumulates,
stores and later releases heat. Polar bears live in the arctic, one of
the most inhospitable environments, yet they can insulate themselves efficiently with the help of their thick fur and blubber as
well as by burying themselves in compact dens. The related project was designed to take advantage of the insulating properties of
the Earth. Its adaptive envelope tracks the sun seasonally, absorbing as much heat as possible to then be released into the compact
interior. The snow leopard lives at high altitudes in the Himalayas
yet it minimizes heat loss with its thick fur and respiratory system.
The project is a relocatable researcher’s laboratory, which has an
expandable system that opens up to thermally protect researchers
in this extreme region.

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Animal Examples — Heat Dissipation  Animals in hot environments are often hairless to increase heat dissipation. Birds do
not sweat but some species, such as storks, defecate on their featherless legs to increase heat loss via evaporative cooling. Camels
allow their body temperatures to rise during the day, drawing heat
away from the vital organs and storing it in their fatty humps.
The large ears of the African elephant and its hairless body also
increase the potential to dissipate heat. Many desert animals, such
as the tarantula and fennec fox, are nocturnal, spending the hot
days in cool burrows and emerging in the cold night. The deep
beak of the toucan is primarily used for feeding, but also contains a network of blood vessels through which body heat can be
radiated to the environment. Large reptiles such as alligators and
crocodiles may bask in pools of water during the hottest parts of
the day to avoid overheating.

Animals use physiological and behavioral mechanisms to dissipate heat in hot environments. (Photographs courtesy
of: yellow-billed storks, B. Larison; Arabian camel, S. Yeliseev; African elephant, A. Kirschel; fennec fox, T. Parkinson;
toucan, J. Drury; tarantula, M. Hedin; American crocodile, M. Tellez)

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Animal Examples — Heat Conservation  Animals living in
cold environments, such as penguins and walruses, tend to be
larger than their relatives in more temperate environments to
minimize their relative surface area. Layers of insulating fat can
also keep the body warm. Cold-blooded animals have different
approaches to keeping warm. Invertebrates such as butterflies
must bask in the open during the warmest part of the day to heat
up. Snowy owls and penguins also possess fine, downy feathers
that can fluff to trap warm air close to the body. Musk oxen and
sea otters have dense underfur for insulation and water-repellent
guard fur to prevent snow or cold water contacting warm skin.
Some fish living in arctic temperatures, such as the Amur bitterling, produce antifreeze-like chemicals that circulate in their
bloodstream and prevent ice crystals from forming.

Animals have many ways to gain or preserve heat, particularily in cold environments. Some can live in extremely
harsh conditions. (Photographs courtesy of: king penguin,; butterfly, S. McCann; walrus, Wikimedia;
snowy owl & sea otter, S. McCann; musk ox, J. Bussey; Amur bitterling, S. Yeliseev)

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10 4

Animal Examples  Reptiles have several behavioral thermoregulatory mechanisms. Collared lizards prefer to keep their
body temperatures at elevated levels and bask in the sun in open
areas to gain heat. On very hot days they raise themselves off the
ground to prevent gaining heat from radiation. Gopher tortoises
dig several very long burrows underground to protect themselves
from heat, losing moisture and predators. They spend most of
their lives in these burrows. Like collared lizards, eastern fence
lizards are often found in open habitats in the sun. They are active
predators and keep body temperatures at an optimum for locomotion and digestion. Sidewinder rattlesnakes get their name from
the way they move across the desert sand. They change from
being active during the day in the cooler months to being nocturnal in the hottest months.

Many reptiles inhabit deserts. Their activity patterns can be modified to prevent them from overheating. (Photographs
courtesy of: collared lizard & sidewinder rattlesnake, P. Niewiarowski; gopher tortoise, S. McCann; eastern fence
lizard, D. McShaffrey)

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10 5

Side-blotched lizard

Uta stansburiana


Chordata Family:
Sauropsida Genus:
Squamata Species:

U. stansburiana

Photograph courtesy of D. McShaffrey

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10 6

Habitat & Climate  Lizards in the genus Uta are known as sideblotched lizards and are common inhabitants of arid regions in the
western United States. There are several species of Uta. The common side-blotched lizard, Uta stansburiana, occurs south to north
from central Mexico to Washington and east to west from western Texas to the Pacific coast. The species prefers open habitats in
deserts and scrublands with elevations ranging from 0 to 2500 m
(0–8200 ft). The project’s location, the portion of the Great Basin
Desert within Utah, is characterized by its northern latitude and
high elevations of 900–2000 m (2953–6560 ft), making it cooler
than other deserts with temperatures ranging from −18°C (−0.4°F)
in January to 50°C (122°F) in July. It rains mostly during the
months of November and April, with an average annual precipitation of 2.5 cm (1 in.). High winds occur during the whole year but
become stronger during the late winter and early spring.


The topography of the Great Basin Desert includes flat valleys interspersed with small mountain ranges. Rocks provide places for basking in the sun as well as places to hide from numerous predators. Sagebrush plants are common
and provide shade in the hot hours of the day. Additionally, the desert basin with eroded terraced cliff sides provides
shaded overhangs. (Photographs courtesy of D. McShaffrey)

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Animal Physiological, Behavioral & Anatomical Elements 
Lizards are ectotherms, meaning they use their environment to
regulate their body temperature. Temperature regulation in lizards is achieved through a combination of their skin characteristics and their behavior. The common side-blotched lizard has a
skin coloration pattern that is typically a dark color on the back
for sunlight absorption and a light color on the abdomen to reflect
heat from the ground.
Lizards obtain or dissipate heat from the environment through
their behaviors. A lizard will adjust its body position to be perpendicular to the sunlight for heat absorption or parallel to the
sunlight while curling up its toes to avoid heat gain by minimizing the area of the body touching the ground. They spend the hottest hours of the day in the shade to prevent overheating.
Large jaw scales,
limited movement

Perpendicular to sunlight
for heat absorption

Parallel to sunlight
to avoid heat gain

Ear scales,

Gular fold scales
small, high range
of movement

Dark color on back for absorption

Hiding in shade
to avoid heat gain

Curls its toes upward
to minimize ground contact

The side-blotched lizard has many behaviors that help its ability to thermoregulate in the desert climate: standing
perpendicular to the sun to absorb heat, standing parallel to the sun to avoid its rays, curling its toes upwards to
minimize contact with the hot floor, and moving to shade when it is too hot. (Photograph courtesy of D. McShaffrey;
project team: J.M. San Pedro, A. Nahmgoong, & Y. Yuan)

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10 8

Interface between the Skin & External World  Lizards’ skin
scales are important for camouflage, preventing desiccation, and
protection against sharp rocks and foliage. The skin consists of
two principal layers: the epidermis, the outer layer, and the dermis, the inner layer. Lizards’ scales are thickenings of the epidermis and are primarily made of a horny substance called keratin,
much like human fingernails.
The scales are thickenings of keratin, connected by hinges of thin
keratin; they are often folded and overlap each other. Areas of
the body with smaller movement have smaller scales to allow for
increased flexibility; large scales are found on areas of the body
with restricted movement. The outer skin is molted periodically
and then renewed by cells in the inner skin layer. Molting allows
room for growth and at the same time replaces worn-out skin.

Horny Layer



Plates of bone
for reinforcement
of horny epidermal

Pigment cell

tissue, blood
vessels and

Lizard scales are one continuous keratin surface formed from the epidermis; they consist of thickened regions (the
horny layer) and thin, connecting hinges. The dermis contains osteoderms and pigment cells (melanophores). The
different sized scales contribute greatly to the lizard’s mobility and its ability to thermoregulate. (Photograph courtesy
of D. McShaffrey)

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Interrelationship between the Skin & Internal Systems  The
common side-blotched lizard’s physiology is optimized for efficiency in extreme temperatures and in water-scarce conditions.
They are opportunistic feeders, readily feeding on invertebrates,
such as mealworms, arachnids or insects, living in their habitat.
Their digestive system absorbs all available water from their prey.
The excrement they produce is a dry white paste, which also
serves as a water conservation strategy.
In addition to aiding with thermoregulation and camouflage,
throat coloration in U. stansburiana reflects a type of mating
strategy observed in males. Males have orange, blue or yellow
throats which correspond to their level of aggression and ability
to maintain pair bonds with females.

Direct sunlight

Water loss
from breathing

heat gain

Thermal radiation
from vegetation

Thermal radiation
from ground

Thermal conduction
to and from ground

The common side-blotched lizard faces many different sources of heat in the desert. Despite these sources, it has
adapted a physiology and behavioral repertoire that allows it to thrive in hot and dry environments. (Photograph
courtesy of D. McShaffrey)

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Proto-Architectural Project  Thermoregulation requires constant vigilance for desert-dwelling organisms, particularly ectotherms. The project investigates how a building envelope system
could respond to the extreme temperature variations of the desert
in order to keep the interior temperature within human comfort
levels. The project is a small artist’s retreat located in the Great
Basin Desert inspired by U. stansburiana’s physiological and
behavioral adaptations to the desert temperature regime.
The design’s focus is to create an enclosure system which maintains 24 hour comfort for the residence during hot, arid days and
sometimes very cold nights. The physiological strategies of the
lizard’s skin have inspired the house walls’ design, while the
lizard’s behavioral adaptations have influenced the “smart” suntracking system actuated by a hydraulic system and sensors.
Great Basin Desert temperature range












Sun-tracking on

Sun-tracking off

Sun-tracking system,
optimal roof angle

Extreme day

Extreme night





75°F–70°F: average indoor temperature

Sun-tracking off

Sun-tracking on



S.C.A.L.E.S. (smart – continuous – active – layered – environmental – system) is the culmination of the observed efficient thermoregulation of the lizard; it combines the characteristics and behaviors that help it survive in the desert,
and integrate it in the building envelope. It takes cues from survival skills of the lizard and makes the building, in
essence, survive in the desert quite comfortably.

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Project Documentation 
S.C.A.L.E.S. uses modular panels,
distributed throughout the entire envelope, in different ways,
depending on their functions. The south-facing wall is composed
of three different types of panels: opaque insulative, photovoltaic,
and operable window. The insulative panel uses phase change
material to allow for a stable interior temperature throughout the
day. The panel is hollow and filled with a bio-based phase change
material. Heat gets stored during the day, while keeping the interior cool. The heat collected during the day is slowly released and
heats the residence at night.
The envelope’s structural system is made up of a braced steel grid
to which the panels attach. All façades follow a similar organizational strategy, while the panel composition may vary depending
on their exposure.

Hot Air


Flexible, waterproof

Cavity for airflow

Phase change material
Cool Air

The photovoltaic panel captures the sun’s rays and converts them to the studio’s electricity. The window panel
allows for views and ventilation. These panels are strategically arranged to maximize their performance. Between
the panels is a flexible, foamed neoprene membrane that allows the panels a range of motion, controlled by sensors,
while being continuous, insulative, and waterproof.

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Converting sunlight
to energy

Accumulates and
releases heat

Providing ventilation
and exterior views

Neoprene/panel joint
(detail on previous
Envelope expansion

Sun-tracking movement

The wall is composed of rhomboidal panels mounted on hardware that allows for a small range of movement. The
scales are mounted on universal joints, gas springs, and rod ends. The universal joints allow for the panel to tilt horizontally and the gas springs move opposite to one another and allow for vertical tilt.

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The artist studio fixes itself directly on the desert floor, much like the lizard. The individual panels gleam in the
sunlight while collecting heat and energy. The structure maintains a slight tilt in the roof and south façade in order
to be optimally positioned to the sun angle.

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Animal Examples  Large ungulates have several adaptations to
allow them to inhabit cold climates. Moose and bison grow dense
fur in the winter to maintain body heat. They also reduce their
metabolism and lower their activity levels to conserve energy
in the winter. Alpacas, along with their thick coats of fur, have
behavioral mechanisms to conserve heat. They protect areas of
their bodies with short fur or thin skin in extreme conditions.
Ermines, or short-tailed weasels, replace their brown summer
coats with a thick white coat of fur in the winter. This coloration
helps to camouflage them in the snow to avoid predation and to
catch prey. They also use dens in the snow to protect them from
the elements.

Many mammals have dense winter coats that allow them to preserve body heat in the winter. (Photographs courtesy
of: moose, B. Miers; American bison, D. Greenfield; alpaca, J. Drury; ermine, L. Parenteau)

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Snow leopard
Panthera uncia


Chordata Family:
Mammalia Genus:
Carnivora Species:

P. uncia

Photograph courtesy of D. Conner, Snow Leopard Trust

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Habitat & Climate   There are five species in the genus
Panthera, commonly referred to as the “big cats.” Snow leopards
inhabit the mountainous regions of central and southern Asia,
their range covering an area of nearly 2 million km2 (772,205
mi2). Snow leopards usually reside at altitudes between 3000 and
5400 m (~9842–17,717 ft). Here, the environment is harsh and
forbidding, with temperatures ranging from −30 to 15ºC (–22 to
The climate is cold and dry with only small amounts of precipitation during the summer months. As a result, the mountain slopes
are sparsely vegetated. Snow leopards prefer these rocky slopes as
they provide a good overview and cover to help them sneak up on
prey. The harshness and remoteness of these habitats means that
it is difficult to study snow leopards in their natural environments.

The snow leopard’s range in the rugged mountainous regions of central Asia stretches through twelve countries:
Afghanistan, Bhutan, China, India, Kazakhstan, the Kyrgyz Republic, Mongolia, Nepal, Pakistan, Russia, Tajikistan,
and Uzbekistan. The home range of a single animal varies from 35 to 1000 km2, depending on prey availability.
(Photographs courtesy of: snow-covered peaks, Snow Leopard Trust; top image by B. Hogue; landscape and leopard,
Felidae Conservation Fund)

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Animal Physiological, Behavioral & Anatomical Elements 
Adults measure around 60 cm (~2 ft) at the shoulders, 1.8–2.3 m
(~6–7.5 ft) between the nose and the tip of the tail and weigh
35–55 kg (~77–121 lbs.). Males are generally 1/3 larger than
females. Most features of the snow leopard relate to thermoregulation. Compared to other cats, their bodies are sturdier and
rounder to minimize surface area and prevent heat loss. Their
small, rounded ears are covered in thick fur. Short forelegs, long
hind legs, and powerful chest muscles allow snow leopards to rapidly pursue prey over unstable terrain. An enlarged nasal cavity
and lung capacity compensates for low oxygen levels at high altitudes. Cold, dry inhaled air is warmed and moistened as it passes
over the delicate tissue covering the nasal turbinate bones, which
also collect heat and moisture with every exhaled breath to help
preserve body temperature and retain valuable water.

Furry Tail

Guard Hairs
Rounded Ears

Nasal Cavity

Thick Under Fur
Large Lungs

The snow leopard uses its long, thick tail for balance when pursuing prey over the mountainous terrain. Densely
furred paws allow movement across snow while the stocky body and thick fur coat prevent heat loss and insulate
in subzero temperatures. (Photographs courtesy of: snow leopard body and face, Snow Leopard Trust, face by F.
Polking, body by M. Trykar; paw, Felidae Conservation Fund; project team: J. Hoen & G. Mamoune)

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Interface between the Skin & External World  Snow leopards
are covered with dense, insulating fur. Hairs on the belly can be
up to 12 cm long. The coat is a white color with brown or graybrown spots during winter, darkening to a light gray color with
dark brown to black spots during summer. These colors provide
superb camouflage all year round. Spots are mainly located on
the animal’s back and are formed by straight guard hairs, which
are up to 70 mm long and between 40 and 60 µm in diameter.
The latter is composed of soft and slightly curved, nonmedullated hairs with a thickness between 5 and 20 µm. This fine layer
of hairs provides most of the thermoregulating properties of the
snow leopard’s fur. The thick tail, which is as long as the body, is
used primarily as a counter-balance during climbing and chasing.
However, it can also be used as a muffler to cover the face when







Microscopic images of the cuticlar structure of the fine fibers and a transverse section of an intermediate fiber. The
snow leopard does not sweat through the skin, which means the thermoregulating properties of the fur are solely of
an insulating nature. Cross-section of skin: 1. Guard hair, 2. Temperature and pain receptors, 3. Hair erector muscle,
4. Epidermis, 5. Dermis, 6. Hair follicle (Photographs courtesy of: leopard cub, Snow Leopard Trust, by H. Freeman;
hair micrographs, A. Galatik)

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Interrelationship between the Skin & Internal Systems  While
fur provides a layer of insulation for the skin, cold air still enters
the body via the lungs due to the necessary act of breathing. The
lungs of the snow leopard are enlarged relative to those of a similarly sized cat in order to maximize oxygen uptake from the thin,
high altitude air. Cold air would chill the body and dry the lungs,
however, reducing overall respiratory abilities.
The nasal turbinate bones housed inside the snow leopard’s nasal
cavity are covered in moist, highly vascularized tissue called
respiratory epithelium. This surface warms and moistens inhaled
air passing to the lungs and reduces cooling of core temperature. The turbinate surface cools as cold air is inhaled but can
trap excess heat and moisture from exhaled air, recouping some
energy and reducing wasted water.










In order to live in extreme altitudes, snow leopards have developed large, powerful lungs capable of extracting
enough oxygen for an active life in a thinly aired environment. The nasal chamber (top image) acts as a two-way air
conditioning unit. Anatomical diagram (bottom): 1. Cranial lobe of the lung 2. Middle lobe of the lung 3. Caudal lobe
of the lung 4. Diaphragm 5. Heart 6. Hyoid apparatus 7. Larynx 8. Trachea 9. Esophagus 10. Scapula 11. Humerus.
Bottom diagram redrawn from Atlas of Veterinary Clinical Anatomy, 2004)

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Proto-Architectural Project  Little research has been achieved
on the snow leopard in its natural habitat due to political problems,
steep elevation and extreme weather. This project is intended to
make their habitats more accessible to researchers through the
design of a structural envelope that could be flown in and deployed
on mountainsides. This structure serves as a base and shelter
for researchers wanting to perform field studies of the cats. The
main responsibility of the building envelope is thermoregulation,
mainly protecting the interior from exterior temperatures varying
from −30 to 15°C (~ −22 to 59°F). Protection from cold, harsh
weather is important for survival in these mountainous regions,
and will only grow in importance worldwide as global warming
causes considerable climatic changes. The snow leopard’s thick
fur, body shape, and respiratory system are prime examples demonstrating nature’s ability to deal with a cold environment.





A: Main structure insulated by vacuumed insulated panels. B: Structure forming the wire frame of a geodesic dome
and made expandable by telescopic members. C: Clusters of rods fixed to each other and a base causing them
to open and close as the skeleton underneath expands or contracts. D: Dark colored, waterproof membrane. By
emulating muscular features of the snow leopard, the proposed structure expands or contracts to create insulating
pockets within the envelope.

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Project Documentation  The multilayered envelope emulates the
anatomical interactions between the animal’s systems. The inner
static structure is based on a geodesic semi-sphere with highly
efficient vacuumed insulated panels, which function as the innermost layer of insulation — expanding and contracting as the frame
moves — and a main structure for the rest of the system to build
upon. A system of telescopic rods, mounted on the inner body,
forms a geodesic dome that pneumatically expands and contracts.
The structure’s connection points contain a round platform hosting a number of hinged and pivoting rods, connecting one end to
the platform and the other end to a corresponding rod. When the
telescopic structure expands, the platforms move apart, forcing the
rods to flatten and open each cluster. A final dark-colored waterproof membrane stretched over the end of the rods unfolds, heating
the enlarged air pocket underneath through solar heating.

Taking cues from the erector muscles in the snow leopard’s skin, the expansion and contraction of the geodesic
structure cause each cluster of rods to open or close according to external climatic conditions. This happens by connecting each rod to a corresponding rod in an adjacent cluster. When the distance between the cluster origin points
increases each cluster is opened.

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12 2


Telescopic Skeleton
Inner Structure

Inflatable Cushions

Close up of façade. An axonometric view of the envelope depicting the geodesic structure and the platforms forming
each cluster of rods. In a typical section through the building envelope the movement in the envelope is ensured by
a series of interconnected inflatable cushions which eliminate the need for local mechanical actuators in each node.

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12 3

The project’s formal expression, shaped by the tensioned membranes, inserts itself within the skyline of the mountain peaks present in the surroundings in which the research facility is located. The pod is a rather curious object in
this environment with its repetitive unified elements and materiality.

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Animal Examples  The cold provides formidable challenges for
mammals. Seals and other marine mammals must endure low
temperatures almost constantly. Large body masses, combined
with a layer of insulating fat called blubber, help maintain warm
core temperatures.
For terrestrial mammals, such as hares, a layer of thick fur helps
insulate the body. Mammals living in extremely cold environments may evolve additional adaptations to the cold. Arctic foxes
have short ears and limbs to reduce their body surface area and
minimize heat loss. A white coat additionally provides camouflage in the snow. The Canadian lynx hunts on deep snow. Broad,
fur-covered feet act as snow shoes, allowing the lynx to run over
the loose snow surface.

Some adaptations that allow animals to thrive in cold conditions are thick fur, a layer of blubber, and reduction
of body surface area. (Photographs courtesy of: elephant seal, J. Himes; arctic hare, A. Holmberg; arctic fox, T.S.
Bortne; Canada lynx, K. Williams)

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12 5

Polar bear
Ursus maritimus


Chordata Family:
Mammalia Genus:
Carnivora Species:

U. maritimus

Photograph courtesy of M. Johnson

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Habitat & Climate  The family Ursidae consists of eight living
species. The polar bear is a particularly distinct species, which
is found throughout the Arctic Circle, across North America,
Europe and Asia. Polar bears are often characterized as marine
mammals as they are comfortable in and out of water and are
dependent on ice as a platform for hunting. The focal region for
this project is the Hudson Bay in Canada.
In this region the average air temperature in summer is 10°C
(50°F) and the minimum winter temperature reaches −23.5°C
(−10°F). Winters are long and dark, with a maximum sun angle
of 6° while summers have long days with a sun angle that reaches
56°. Some arctic areas are covered with ice all year long, while
others may lose ice, allowing the arctic tundra vegetation to



Polar bear distribution includes the U.S. (Alaska), Canada, Russia, Denmark (Greenland), and Norway. Polar bears
spend most of their lives on ice. Their range spans the arctic regions of all continents where they roam pack ice
searching for seals. They venture on land only during the summer, when the ice melts. (Photographs courtesy of:
sunset and polar bear, M. Johnson; glaciers, G. Rochette,

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Animal Physiological, Behavioral & Anatomical Element 
Polar bears are extremely large. Males can measure up to
3 m (~10 ft) in length and weigh an average of 500 kg (1100 lbs).
Females may weigh half as much as males. The large body mass,
combined with short limbs and ears, reduces the surface area to
volume ratio of the body preventing loss of body heat.
Thick fur and a layer of insulating fat provide the primary means
of insulation against the harsh arctic cold. Polar bears hibernate
through the winter in dens dug into snow banks or the ground.
Polar bears roam vast distances over the sea ice in summer
searching for seals. Large, broad paws and sharp claws provide
snowshoe-like assistance when walking over snow. They are
excellent swimmers and have been observed to stay in the water
for prolonged periods of time when moving between ice floes.



exchanges heat
& moisture


minimizes heat loss

black to absorb
the heat of the sun




over 10 cm of fat


provide grip for icy


long guard hairs &
short wooly hairs


enable stability,
balance & swimming

The polar bear is the largest living carnivore. Its entire appearance reflects life in the cold, hostile arctic. Newborns
are less well protected and are born in dens dug into snow or earth. They emerge in summer, already able to tolerate subfreezing temperatures. (Photographs courtesy of: snout and mother with cub, M. Johnson; project team: im
studio mi/la & I. Mazzoleni)

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Interface between the Skin & External World  Arctic temperatures can reach −45°C (−49°F) during the depths of winter.
The polar bear’s primary defense against the cold is its skin.
Externally, polar bears are insulated by a thick, white fur coat
that covers almost the entire body, including the soles of the feet.
The fur is composed of two layers. The dense white underfur provides the main source of insulation by trapping warm air close to
the skin and, conversely, by preventing contact by ice and water.
In the outer layer, longer guard hairs are hollow and transparent, lacking pigments which also act as camouflage in the snow.
Polar bears molt during the summer. Beneath the skin within the
hypodermis, polar bears are further insulated by a layer of fat.
Additional features, such as black skin, particularly on exposed
areas such as the nose and tongue, allow the bears to absorb solar
energy and prevent additional passive heat loss.

takes in water when
swimming, acting like a
wet suit
sun rays

traps heat close to the

maximizes the absorption
of solar radiation


insulates bear from low

striated & highly

Fur is the polar bear’s primary defense from the harsh cold. The dense underfur traps warm air close to the skin,
warming the body. Longer, hollow guard hairs protect the underfur from the elements. The skin is also effective in
providing protective layers that are capable of trapping water as the bear swims in order to reduce heat loss. Beneath
the skin, a thick layer of fat provides additional insulation. It has been proposed that the hollow hairs guide UV radiation to the skin, similar to fiber optic cables, but this now seems unlikely. (Photograph courtesy of D. McShaffrey)

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Interrelationship between the Skin & Internal Systems  Like
all mammals, polar bears are endothermic, meaning that they
produce their own body heat internally. Their large body mass,
combined with short extremities, thick insulating blubber, and
fur, make the polar bear exceptionally efficient at retaining body
heat. In fact, overheating becomes problematic for polar bears,
even in subzero temperatures. This prevents polar bears from
undergoing prolonged running. Swimming or rubbing themselves
in snow are efficient ways of cooling down.
Viewed through an infra-red camera, polar bears are almost
invisible; only the warm air leaving the nose and superficial blood
flow to the less furred head provide signs of life. Also, the blood
cells are adapted to hold more air, allowing for longer stints under





The polar bear’s large body mass reduces its surface area to volume ratio, which helps to retain body heat. With thick
fur, polar bears are so efficient at retaining heat that they appear almost invisible to infra-red cameras; only warm
air lost via the nose can be distinguished. (Photograph courtesy of D. McShaffrey; anatomy diagram adapted from
National Bowhunters Education Foundation [2006]; thermal diagrams adapted from P. B. Dill and L. Irving [1964])

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Proto-Architectural Project  This project mimics the physiological and behavioral adaptations which have allowed the polar
bear to survive in the planet’s harshest weather conditions. The
project attempts to optimize natural local resources to provide
energy needed to regulate human thermal comfort. The compact
living units are partially embedded in the earth, similar to the
bear’s hibernation den. The units’ southwest orientation optimizes the heat gain from the sun. The sun’s energy, heat and light,
is harvested by the active building envelope composed of hollow
re-orientable fur-like glass tubes and travel to the insulating strata
where the energy is stored, conserved and slowly released into the
compact pod. The phosphorescent cells embedded in the phase
changing material (PCM) collect light, which is slowly released at
night creating an atmospheric sky-like vaulted ceiling.

Studies of optimized shapes were facilitated by computer modeling to aid in understanding and generating complex
geometries. Computational/digital studies were further developed through testing with rapid prototyping and by
engaging three-dimensional drawings to further explore the inherent spatiality. These preliminary investigations are
necessary to proceed in the development and fabrication of full-scale mockups to test the feasibility of current ideas
and material strategies.

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Project Documentation  The envelope is a storage for heat and
light which minimizes the use of active systems and maximizes
the use of passive systems. The storage of heat and light within
the active envelope varies according to seasonal changes. During
the winter, light and heat are transferred into the unit, where heat
is conserved within building-integrated thermal storage in the
form of PCM. During the long, cold summer nights, light is managed and minimized while the heat is still being collected. This
variability in conveyance and conservation of heat and light is
accomplished via sensors within the movable tubes which track
the sun angle and optimize the heat accumulation. The thick
envelope assembly controls a number of crucial factors, including
thermo-physical properties of the materials, the outdoor climate,
and the operating schedule of the compact dwelling.



air vent
heat / light
thermal mass



light shaft






thermal mass

The dwelling is articulated around a compact central space to avoid energy dispersion. Each area satisfies multiple
programmatic functions. For example, the lounging area is the living space as well as the sleeping space, while the
steps allow for small gatherings to observe the sky by looking through the scattered openings on the ceiling.

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This structure is a super-efficient habitable cell. Its openings, diffusely directed, capture all the possible energy from
the sun, accumulating light and heat and dispersing them through high-tech materials and technologies. In exploiting
the available energy of the sun, this unit is a comfortable shelter, welcoming life and providing protection from the
relentless environment of the extreme north.

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The cold that penetrates the skin deeply inside. The reverberation of the light on the white flat polar
landscape, so strong, almost unbearable. The horizon so far, feels endless. The lost, bewildered look
expresses the void. All of a sudden, something is delineated, something starts to reveal itself: the vision, the
imagination of something, perhaps a glare, some shapes, undefined creatures, feral structures.

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  Water Balance


Amphibians such as frogs have a
close relationship with water. Their
skin is permeable, and many stay
moist by living in or near water.
(Photograph courtesy of S. McCann)

Life on Earth has an intimate relationship with water. All organisms, from humans to bacteria, are primarily composed of water;
it is the main component of our cells, and most of our tissues and
organs are bathed in it. It is through water that nutrients, vitamins
and hormones are delivered to different parts of the body where
they are used for producing energy and carrying out life’s basic
functions. Different environments place strong demands on water
use and, as such, animals exhibit many diverse adaptations for
collection and conservation of water and protection from being
exposed to too much water. On one extreme, organisms in harsh
environments like the desert struggle to acquire enough water to
survive, while on the other extreme aquatic organisms are surrounded by water and must prevent their tissues from becoming
overcome by it. While different organisms have different relationships with water, maintaining a balance of water within the body
is crucial for survival.
Water cannot be wasted in the desert environment, because it
can only be acquired from a few sources. Most desert creatures
obtain water from plants, particularly those that are adapted to
store water, like cacti. Others meet all of their water needs from
the food they eat. Desert-adapted animals have evolved innovative strategies to gain water and prevent water loss. Some retain
water in fatty tissue within the body; when the fat is metabolized
the water in the tissue is released and can be used by the animal. Reptiles and birds excrete waste in the form of uric acid,
which wastes less water than the urea excreted by amphibians

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and mammals. Many reptiles excrete very dry feces to further
preserve water. Desert-dwelling mammals such as kangaroo rats
possess highly efficient kidneys and excrete extremely concentrated urine to retain ingested water. Exhaled water vapor may
also be recaptured in the nasal cavity to prevent further water
loss. Behavioral adaptations such as burrowing underground not
only provide a cooler temperature in the day, but can also aid in
water balance, because soil is cool and moist.

Animals with permeable skin use
many mechanisms to prevent the
loss of water from their tissues. For
example, snails and salamanders
both secrete mucus in order to stay
moist. (Photographs courtesy of:
snail, P. Thompson; salamander, H.

11_WaterBalance.indd 136

Though not as extreme as desert conditions, other biomes possess their own set of water conservation challenges that must be
overcome by the organisms living there. In all terrestrial habitats
the concentration of water in the environment is lower than it is
inside the body of animals. Many animals have thick outer skin
layers that prevent them from losing water to the environment.
Arthropods generally have a waterproof wax within the top layer
of their exoskeleton; they can also close pores used for breathing to prevent additional water loss in dry conditions. Reptiles
and mammals use the protein keratin in their skin as a barrier to
water loss. Mammals may also produce an oily film to cover and
waterproof their skin and fur. The feathers of birds lock together
and align in such a way as to confer waterproofing.15 All cells use
salt ions, which are charged particles, to function. As the ions
flow in and out of cells, they create electrical gradients across the
cell membranes. These gradients are integral to all aspects of cell
function, from transporting sugars for energy, to firing of neurons
that transmit information from the brain to muscles. Maintenance
of the concentration of salts to water, or osmoregulation, is critical
for most animals. Animals that live in or have a close association
with water do not have an issue with preventing water loss; however, they do need to keep the concentration of molecules in water
and the concentration of molecules inside their bodies balanced.16
Architecture has always been concerned with issues related
to humidity control and moisture penetration, and a lot can be
learned from nature when it comes to water management. Water
balance in wet climates has traditionally focused on keeping out
rain and humidity, while in arid climates architects have learned
to add humidity to dry air. In traditional Islamic architecture, for
example, the insertion of small fountains in courtyards provides
comfort by means of evaporative cooling. In tropical climates
strategies to achieve comfort are achieved by optimizing natural ventilation and detachment of the structure from moist soil.

13/12/12 11:40 AM


The Alhambra region is rich in fountains and courtyards for evaporative
cooling to cool the rooms surrounding the patio. Patio de los Arrayanes,
Alhambra, Granada, mid 14th century. (Photograph courtesy of L. Pretorius)

Today, water conservation has become a fundamental driver of
design innovation throughout the world in order to lessen our
environmental impacts. Increasing population sizes have led
to the establishment of new settlements in areas of the planet
where humans struggle to find the basic resources required for
sustainability, such as water. Strategies for water conservation
can be developed at many design scales, ranging from low-flow
plumbing fixtures to irrigation systems, and include both collection in cisterns and the reuse and treatment of gray and black

Court of la Acequia. Palacio de
Generalife, Alhambra, Granada,
1309. (Photograph courtesy of L.

11_WaterBalance.indd 137

An understanding of water balance in the banana slug, with its
permeable, mucus coated skin, led to the design of a greenhouse
that collects water in manmade bladders (or cushions) that provide irrigation to plants. The tropical environment of the dyeing
dart frog inspired the development of museum walls designed to
remove moisture from the air to protect valuable artifacts. The
ochre sea star’s ability to live both under and above water motivated the design of a retractable structure growing under a pier.
Namib desert beetles’ adaptations to extremely harsh environments inspired the design of a research facility that, by capturing
fog, provides water for human use.

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Animal Examples — Water Balance  Marine invertebrates such
as sea slugs, jelly fish and anemones possess thin body walls that
allow for diffusion of gases and nutrients via osmosis. These animals maintain a salt concentration in their tissues that is the same
as that of the surrounding seawater in order to balance the concentration of salt to water in their cells. On land, different strategies
are required to prevent water loss. Crabs spend part of their time
out of water and possess tough exoskeletons that prevent desiccation. Completely terrestrial invertebrates such as earthworms that
lack an exoskeleton are restricted to moist environments to prevent
loss of water. Most insects possess an exoskeleton that frees them
from this constraint. However, the exoskeletons of the smallest
insects, such as springtails, are so thin that they cannot prevent
water loss. These tiny animals are therefore restricted to moist
environments where the risk of passive water loss is minimized.

Animals that live outside of water have several mechanisms to prevent desiccation. Animals that live in water still
have to regulate the balance of salts and water in their cells. (Photographs courtesy of: sea cucumber, A. Jaffe;
tadpole, K. Pease; octopus, B. Larison; earthworm,; jellyfish, J. Himes; crab, J. Drury; springtail, S.

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Animal Examples — Water Balance  Marine fishes such as
eels maintain a lower salt concentration in their tissues than in
the surrounding water by actively regulating salt levels through
the gills and by excreting concentrated urine. Amphibians retain
many similarities to their fish ancestors, as indicated by the fully
aquatic tadpole stage. Adult amphibians retain thin skins, allowing some gas diffusion that makes them susceptible to dessication. Thus, amphibians are restricted to semi-aquatic habitats, as
found in newts or caecilians burrowing in moist soils. Whales
and dolphins are mammals that have returned to the water – these
animals face similar challenges as marine fishes and have a thick,
nonporous skin and efficient kidneys for maintaining low blood
salt levels. Fetuses are bathed in fluid which provides cushioning
to the developing animal, transports nutrients from the mother
to the baby, and is the medium through which breathing occurs.

Animals that live outside of water have several mechanisms to prevent desiccation. Animals that live in water still
have to regulate the balance of salts and water in their cells. (Photographs courtesy of: salamander, A. Illum; eel, A.
Jaffe; pangolin fetus, K. Benirschke; frog & dolphin, H. Thomssen; Caecilians, J. Measey; anemone, J. Himes)

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14 0

Animal Examples  Many invertebrates have soft, permeable
skin, making them vulnerable to desiccation. Consequently,
they often live in moist environments, either in or near water.
Turbellaria are a group of mostly free-living flatworms with ribbon-like bodies. Turbellaria have to balance the concentration of
salts and water within their bodies, achieved through a specialized system of cells called protonephridia. Land snails keep their
skin moist by producing large quantities of mucus and retreating
into their shell, which has an opening that can be plugged with a
thick layer of mucus while inside. Leeches are segmented worms
that primarily live in freshwater habitats such as streams to maintain continuous moisture. Isopods are crustaceans, but unlike
other crustaceans they do not live in water. Many isopods have
evolved to live in humid habitats, as they breathe through gills
that must remain moist.

Invertebrates have physiological adaptations to prevent water loss. Turbellaria and land snails have mucus-secreting
glands for protection and to stay moist. Isopods excrete waste in a dry form, which helps to conserve water.
(Photographs courtesy of: turbellaria, Wikimedia, H. Krisp; land snail, J. Robinson; leech, Wikimedia, C. Schuster;
isopod, Wikimedia, G. San Martin)

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Banana slug

Ariolimax columbianus


Mollusca Family:
Gastropoda Genus:
Geophila Species:

A. columbianus

Photograph courtesy of D. McShaffrey

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Habitat & Climate  There are three species of banana slug:
Ariolimax californicus, A. columbianus and A. dolichophallu.
The California banana slug, A. californicus, is the focus of this
project and lives on the forest floors of the Pacific coastal rainforest belt, in foggy and forested habitats. The climate of this
area is regulated by its proximity to the Pacific Ocean, resulting in a Mediterranean like climate. As a result, the average high
temperature ranges from around 15°C (59°F) in winter to 22°C
(~71.5 °F) during the summer months.
Average annual precipitation is around 495 mm (19.5 in), with
most rainfall occurring between November and April, while
little or no precipitation falls during the summer months. There
is an annual average of 70 days with measurable precipitation.
Summers in the Monterey Bay area are generally cool and foggy.



The banana slug can be found on the floors of the Pacific coastal rainforest belt from southeastern Alaska to Santa
Cruz, California, where the project is located. Banana slugs are found in relatively mild and humid climates due to
their thin, permeable skin. Due to the need to remain moist at all times, banana slug activity is primarily nocturnal.
(Photographs courtesy of: habitat, P. Guerrero; banana slug, D. Siciliano)

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Animal Physiological, Behavioral & Anatomical Elements 
Slugs are most active after rain because of the wet ground, while
during dry summer conditions they cover themselves in damp
areas under fallen logs or rocks. Breathing occurs through a pallial lung — a heavily vascularized tissue of the dorsal body wall,
or mantle, that allows for gas exchange. The pneumostome is an
opening in the mantle that lets air into the mantle cavity/lung. The
rate at which it opens and closes relates to external temperatures,
metabolic rates and hydration levels. Slugs also breathe through
their skin, which allows for passive gas exchange between the
slug and the atmosphere. On the head of the slug are four tentacles used to sense its surroundings: the two upper optic tentacles
observe light and movement, while the two lower sensory tentacles discern chemicals. The slug’s mouth is located at the base
of the head, where its file-like tongue is used to scrape up food.




Vascular network
connected with


The banana slug contains a pneumostome to allow for gas exchange. Inside the mantle cavity the slug has a highly
vascularized section of tissue that facilitates gas exchange. The vascular network diagram is adapted from a diagram of a snail, assumed to be similar. (Diagram adapted from T.R. Jones; general anatomy diagram adapted from
Wikimedia & R.A. Barr [1927]; project team; A. Bang, M. Alam: & J.S. Pedersen)

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14 4

Interface between the Skin & External World  Banana slugs
live in humid environments to ensure moist skin. They also
secrete mucus to prevent desiccation, aid in respiration across a
moist surface, find and attract mates, and aid in movement, among
other functions. Slime deters predators by its consistency but also
because it has anesthetic, or numbing, properties.
During periods of unfavorable weather, such as hot spells or cold
temperatures, the banana slug estivates, covering itself under
leaf litter and secreting a layer of mucus around its body. Banana
slugs are named for their characteristic yellow coloration. They
often have golden yellow bodies with dark spots; however, they do
exhibit variation in color and can be brown, black and sometimes
white. These color patterns can vary with changes in diet, moisture, light availability, age and health.





The foot is the largest part of the body, consisting of many tiny muscles that help the slug crawl about in wavelike
motion. A microscopic view of banana slug mucus granules; these granules are broken open and mucus is released.
(Photographs courtesy of: slug close-ups, D. Siciliano; internal anatomy diagram adapted from G.M. Barker [1999];
mucus granules, Luehtel et al. [1991])

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Interrelationship between the Skin & Internal Systems  Slugs
produce multiple types of mucus, ranging from thin and watery
to thick and gooey, which serve different functions. Thick mucus
protects the slug’s body from the surface of the ground, prevents
dessication, and helps in locomotion. Together with the foot’s
muscular contractions, this type of mucus helps the slug move
forward. Another layer of mucus is thin and watery and is spread
from the center of the foot to the edges of the foot. The mucus is
made up of fibers that prevent the slug from slipping down vertical surfaces. It also has thixotropic qualities, meaning that when
the slug is standing still, the mucus sets into a comparatively firm
mass, but as soon it starts moving the mucus liquefies. Mucus is
hygroscopic, which means that it rapidly attracts water molecules
from the surrounding environment. Dry mucus is contained in
granules, and when broken open mucus is released as necessary.



Mucus for

Mucus for


The protective mucus is secreted from the top of the slugs back and then distributed over the body. The mucus that
helps the slug move forward is secreted from tiny glands in front of the foot and acts as a track that the slug can
glide along. (Photographs courtesy of: standing, D. Siciliano; mucus trail, P. Guerrero; diagram adapted from R.A.
Barr [1927])

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14 6

Proto-Architectural Project  The program of the building is a
greenhouse placed in humid Santa Cruz where people can come
to observe and learn about forest ecosystems. Four characteristics
of the slug were initially analyzed and explored in relation to the
design of building: 1) the skin’s porosity and permeability which
enable breathing; 2) the mucus’ ability to protect the slug against
both dessication and predators; 3) the slug’s ability to adapt to
changes in humidity and temperature in relation to the surrounding environment; 4) the slug’s ability to maintain balance and
communication with its environment through homeostasis. The
greenhouse has an adaptive envelope which adjusts and changes
according to weather conditions. The envelope fluctuates, changing the internal environment by allowing sun, air and rainwater to
permeate inside when desired, and shielding the vegetation from
the elements in other instances.

The greenhouse has both an irregular elliptical shape and foundations, thus adapting to the irregularity of the tree
canopies above and to the organic uneven forest floor. The process of rainwater collection — later used for plant
irrigation – changes the performance, look and feel of the envelope.

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Project Documentation  The structure is a steel grid which
holds the aggregation of bladders, or cushions, in place. The
majority of the cushions are composed of a two-layered silicone
structure filled with water, whereas the roof cushions are only one
layer to collect rainwater, exposed to the outside and allowing the
rainwater to drip straight into the structure, enabling transport
through the envelope. This transport mainly happens through a
secondary structure of woven plastic tubes within the cushions.
As the water fluctuates the bladders inflate or deflate, forcing
the envelope to close or open accordingly. During rainfall, the
silicone units fill, get heavier, and start to droop downwards and
open up the wall as they tug the clamps holding them in place.
This allows rain to flow into the greenhouse to irrigate the plants,
while the overflow is stored for further irrigation. As the envelope
dries up and the water dissipates, it will close off the wall.




Shape of the
silicone when empty

Single layer
roof cushion

Roof cushion detail open to collect rainwater. The envelope uses the cushion deformations provoked by the water
to close off or open up according to the moisture in the atmosphere just as the slug uses its mucus to regulate the
degree of air and water exchange through its skin.

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14 8

The wall section provides details of the integrated systems within the envelope. The clamps that fasten the silicone
cushions to the structure are made of a thicker, less elastic silicone, thus making the clamps substantially rigid. The
cushions themselves can stretch and expand according to the quantity of the contained rainwater. The photograph
shows a mock-up testing four cushions.

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The interior of the greenhouse is dominated by droopy water-filled silicone cushions. This interior volume changes
over the day and year. The envelope is active and flexible, registering the ever changing climatic and environmental

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Animal Examples  Amphibians are unique among terrestrial
vertebrates in possessing skin that is permeable to both air and
water. Even though many amphibians are primarily terrestrial,
they live closely associated with water in moist environments.
The black-bellied slender salamander is in a family of salamanders that lack lungs; that means they only respire through their
skin and the tissue inside their mouths. They are most active at
high humidity levels and are found underground or under rocks
and logs when the humidity levels are low. The squirrel treefrog is
also found in moist habitats and hides under bark and leaves when
inactive. Eggs are laid and larvae develop in ponds and swamps.
The granular dart frog has bright warning coloration, and is found
in lowland tropical forests. Adults are ground dwelling, but after
the eggs are laid and develop into tadpoles, the mother brings
them to water-filled bromeliad plants in the tree canopy.

Amphibians primarily obtain water from their environment through passive diffusion through their skin. (Photographs
courtesy of: salamander, M. Hedin; lungless salamander & squirrel treefrog, S. McCann; granular dart frog, J.

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Dyeing dart frog

Dendrobates tinctorius


Chordata Family:
Amphibia Genus:
Anura Species:

D. tinctorius

Photograph courtesy of M. Bartelds

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Habitat & Climate  Poison dart frogs are native to moist environments in Central and South America and are all members of
the Dentrobatidae family. Many have colorful skin patterning
and are toxic. Dendrobates tinctorius, or the dyeing dart frog,
occurs in the eastern Guiana Shield, in southeastern Guiana,
Suriname, French Guiana and northeastern Brazil. They occur in
isolated populations in the uplands of the Guiana Shield, preferring elevations of 200 to 600 m (~650–2000 ft). This species does
not prefer lowland forests, perhaps because they experience seasonal flooding, and the frog is primarily ground dwelling. With
an average of 373 cm (147 in.) of rain per year, precipitation is
highest between April and June, and lowest between August and
September. Yearly relative humidity averages 76.5% during the
summer months, with peak levels in May (83%).



The forests of French Guiana remain moist through much of the year, providing ideal habitat for the dyeing dart frog.
D. tinctorius is primarily ground dwelling and can be found in tree roots, on vines or under rocks, where it forages
in the leaf litter. (Photographs courtesy of M. Bartelds)

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Animal Physiological, Behavioral & Anatomical Elements 
Like other poison dart frogs, D. tinctorius occurs in extremely
humid areas, most often near running water. They are most active
during the rainy season, foraging in canopy-gaps on arthropods.
In the dry season they retreat to protected areas in trees and
plants and consume less prey. They are diurnal, meaning they are
active during the day, particularly in the early morning and late
Unlike many other vertebrates, frogs do not have waterproof skin,
and their lifestyle is very closely associated with water. In fact,
larval development takes place in the water. While lungs function
in gas exchange, frogs breathe primarily through their skin. Their
skin has to remain moist for breathing, and this is done with the
aid of mucus glands in their skin.

Skin + Lungs

The skin aids in thermal regulation, UV protection, warning predators, respiration and toxicity. The pigment melanin
absorbs heat during the day and helps block UV light. Each foot has four toes which possess a pad that functions as
a suction cup used for gripping. (Photograph courtesy of M. Bartelds; project team: E. Lani & J. Su)

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15 4

Interface between the Skin & External World  Frog skin is permeable and moist. Their skin consists of two main layers, the epidermis, or outer layer, and the dermis, the inner layer. Poison dart
frogs are recognized for their bright coloration and toxic skin. The
color patterns serve as a warning to predators; they are a signal of
their toxicity and distastefulness. The skin contains specialized
glands located in the dermis that produce poison. The poisons
are neurotoxins, called batrachotoxin, which act on the nervous
system of the animal that ingests them. Poison is squeezed out of
the glands when muscle cells around the glands contract. Toxicity
is thought to be acquired from the diet, because in captivity frogs
often lose toxicity. Mucus glands are much smaller and produce
mucus to keep the skin moist. This moisture helps prevent desiccation and also serves gas exchange functions. Both types of
glands are invaginations of the epidermis.


Sodium transport
Permeability barriers


Melanin production
Skin lubrication
Toxin production



Skin coloration patterns are produced from pigmented cells, called chromatophores, which are located in the dermis. Because dyeing dart frogs occur on isolated islands, each population possesses striking variation in its skin
coloration patterns. D. tinctorius can be any combination of blue, yellow, white and black with stripes and spots.
(Photograph courtesy of M. Bartelds)

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Interrelationship between the Skin & Internal Systems  Gas
exchange occurs through the lungs and the skin. Oxygen passes
directly through the membranous skin and into an extensive
network of blood vessels. When using their saclike lungs, frogs
breathe through opening their mouth or nostrils and allowing air
to flow through. Once oxygen makes it into the blood stream, the
oxygen-enriched blood enters into the heart, which is made up
of only three chambers, two upper atria (right and left) and one
lower ventricle. Oxygen-enriched blood is forced into the arteries
and disperses to tissues throughout the body, while oxygen-poor
blood is enriched through the skin and lungs and then distributed
accordingly. The drink patch, densely populated capillary tissue
located on the underside of frogs, can be submerged or touched
against a moist surface to hydrate the body.






The drink patch is located between the hind legs on the lower belly, enabling the frog to easily lower it into the
water to rehydrate. The frog simply dips its pelvic area below the water surface or it presses the area against a moist
surface. Water molecules are wicked through the skin by osmotic action and delivered to organs throughout its body
on the cellular level.

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Proto-Architectural Project  The museum, located in Matoury,
is dedicated to Amazonian cultural history. It houses both indigenous and imperial artifacts and art. Lifted off the ground, the
building is designed with a concentric plan, making humidity and
thermal regulation manageable. The more moisture tolerant artifacts are located closer to the perimeter while the more delicate
objects are placed in the center. Using the D. tintorius as its inspiration, this project has led to an architecture that mediates humidity levels from an exterior expanse to interior, intimate spaces.
Density, layering and curved surfaces play a large role in its aesthetic, but most specially it functions to dehumidify the museum
space. Rather than capturing water that is generously available in
the rainforest, it explores how to keep humidity out. The humidity
regulation is mostly passive, but supplemental mechanical systems are included for use in extreme weather.

Zone 2
45% humidity

Zone 4
35% humidity

Zone 1
65% humidity

Zone 3
55% humidity



The design seeks to wick water out of a space by capturing it in a desiccant-like structural system. Walls become
denser and thicker as they approach the center of the symmetrical plan. This removes humidity at the graded percentages and surrounds the most precious artifacts with a large buffer zone, intensifying the experience of depth
and layering within the walls.

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Project Documentation  The first layer is the permeable one, a
hard material with more or less perforation according to the specific humidity desired. The second layer is a paneled network of
tubes filled with chilled water. The cooler temperatures within the
tube attract water through condensation, which is absorbed by the
third layer, a structured foam desiccant. The desiccant maintains its
shape and cannot degrade too readily under moist conditions, with
a cellular anatomy creating space between molecules large enough
for water to pass through. Behind this is a second “selectively” permeable layer, 6, that acts as a surface weep screed, allowing water
to weep from the desiccant into the wall cavity, but preventing the
re-admittance of water into layer 3. Through the main structure and
a secondary tension structure, dry air would flow wicked water to a
designated catch, to be dumped under the waterproof “belly” of the
building, to act as a cooling mechanism.

Top diagram shows how the selectively permeable layer is inspired by the scalar coloration of D. tinctorius’ skin. The
layering of the various wall tiles is necessary to dehumidify but also to create a depth and scale that visitors perceive
as their museum visit progresses.

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coating to prevent
reverse absorption
from humidity
wicking air-flow

Light cable and tension
structure to allow for
maximum strength and


Formed foam

6” 1’-0”

Zone 1




Zone 4

Tar coated structure
w/ vents in blocking for
a closed system
circulating forced air

Chilled tube condensers with
light tubes intermittently

Selectively permeable
layer of pre-formed
panels of a water
resistant material

Exploded wall section illustrating each layer: 1) selective permeable layer; 2) chilled condensers and lighting; 3)
structured foam desiccant; 4 & 5) lightweight post & tension structure; 6) secondary selectively permeable layer.

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The museum spaces surround visitors with their walls composed of capillary-like tiles that facilitate air dehumidi­
fication. Illuminating the room are lights with blue hues, while highlighting artifacts and walls are yellow hues.

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Animal Examples  Marine invertebrate animals, especially
those that live in intertidal zones, have to deal with the pressures of
desiccation as well as variations of water salinity. Chitons inhabit
shallow waters and have eight hard plates covering their bodies.
For protection from these environmental pressures they often live
under rocks or in rock crevices. They quickly regulate water balance in their cells when salt levels are too high or low. Blood sea
stars are found in coastal regions and can be found on or under
rocks to protect themselves from predators and external elements.
Barnacles are sessile crustaceans that primarily live in the intertidal zone. They have shells that help prevent them from desiccation. The strawberry anemone is another sessile animal. However,
they can detach themselves from the substrate when conditions
such as desiccation cause them to move to another location. They
regulate the salt concentration in their tissues through diffusion.

Many marine invertebrates live in harsh environments where they face large variations in water availability and
salinity. (Photographs courtesy of: chiton, R. Muthukrishnan; blood sea star, A. Jaffe; barnacles,;
strawberry anemone, M. Bartosek)

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Ochre sea star

Pisaster ochraceus


Echinodermata Family:
Asteroidea Genus:
Forcipulatida Species:

P. ochraceus

Photograph courtesy of H. Thomssen

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Habitat & Climate  Pisaster ochraceus inhabits rocky shores,
kelp forests and estuaries along the Pacific coast of North
America. It is most common in the rocky intertidal zone, one of
the harshest and most dynamic environments on Earth. Rocky
intertidal organisms are subjected to myriad predators from both
the land and sea, as well as intense and nearly constant wave
action. In addition, they are exposed to air and direct sunlight up
to two times per day at low tide. While P. ochraceus has an extensive range along the Pacific coast, this project focuses on southern
California, with a Mediterranean-type climate of sunny, warm
summers and mild, wet winters. The rainy season (November
to March) rarely produces more than 30 cm (12 in.) of rainfall.
Ocean temperatures range from 13 to 21ºC (56–70ºF), with an
average relative humidity of ~65%. Temperatures rarely exceed
85ºF (29.5ºC) in summer, nor drop below 50ºF (10ºC) in winter.



Pisaster ochraceus is abundant on rocky, intertidal shores along the entire Pacific coast of North America from Alaska
to Baja, California. Animals that live in these harsh habitats must be adapted to survive intense wave action and to
resist desiccation at low tide. This project is focused on the intertidal zone at the Santa Monica pier. (Photographs
courtesy of: beach and tide pool, N. Lucas; pier, R. Molina)

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Animal Physiological, Behavioral & Anatomical Elements 
Pisaster ochraceus can grow to a diameter of 25 cm (10 in.) with
five arms radiating from the center. It is found in two distinct color
morphs, with purple being slightly more common than orange. It
has no centralized nervous system (i.e., brain and spinal cord), but
instead a series of connected nerves that run throughout its body.
A unique physiological trait of sea stars and other echinoderms
is a water vascular system. The system uses a network of canals
and muscles to move water to transport food and waste and for
respiration. It also uses the system to create and relieve suction
among hundreds of tube feet independently, allowing the animal
to “walk” along the substrate. This means that P. ochraceus can
readily move around underwater but must remain largely in place
while exposed at low tide.



The figure above illustrates the internal pneumatic ring canal (circulatory), stomach (digestion), ampulla (circulatory),
and tube feet (locomotion/predation). The ring canal moves water taken in from a dorsal valve (madreporite) and
channels it through the arms of the ochre sea star to the ampulla and finally to the tube feet as it moves in the water.
(Photograph courtesy of D. McShaffrey; project team: P. Mecomber & A. Ariosa)

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16 4












The skin of Pisaster ochraceus is a continuous epidermis with embedded bony plates called ossicles. The ossicles
create a network of armor plates that are locked closely together when out of the water, producing a hard and tough
skin. A strong vacuum is also produced inside its many tiny tube feet, making it nearly impossible for waves or predators to dislodge the animals from their rocky substrate. Evaporation of water through dermal branchiae helps keep
the animal cool while exposed to sunlight and air. (Photograph courtesy of D. McShaffrey; drawing adapted from R.
Fox, Lander University)

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Interrelationship between the Skin & Internal Systems  Living
in the intertidal zone, P. ochraceus must be adept at handling two
entirely disparate environments. Its body is essentially a pneumatic machine pumping water through its water vascular system
to move its limbs. The system branches out to five radial canals,
one for each arm of the sea star.
Desiccation, the loss of water via evaporation, is critical in setting
the upper limits of many intertidal species. However, P. ochraceus
can tolerate a loss of up to thirty percent of its weight in bodily
fluids. It also behaviorally combats desiccation by hiding under
rocks to avoid direct sunlight, but will move about as it searches
for food — particularly for the California mussel, Mytilus californianus, an abundant intertidal organism that has its own unique
adaptations for dealing with the harsh habitat.





When underwater, the ochre sea star becomes softer and more malleable, as soft tissue is exposed to aid in respiration and waste excretion, and water is pumped in and out of the tube feet for locomotion. The skin is also covered
with tiny pinchers called pedicellaria that are used underwater to prevent larvae of other marine invertebrates from
settling and growing on its surface. The ability of P. ochraceus to respond to its dynamic intertidal environment in
these ways is largely made possible by the water vascular system described above. (Photographs courtesy of D.
McShaffrey; drawing adapted from R. Fox, Lander University.)

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Proto-Architectural Project  The project is positioned in the
intertidal region of Santa Monica, California, a natural habitat
for the ochre sea star. The iconic Santa Monica pier, jutting out
hundreds of feet over the ocean, is the perfect place for tourists to
experience this region. The proposal re-interprets both the Santa
Monica pier and the skin of P. ochraceus. The project is designed
under the pier, using the existing infrastructure to place a new spa
and observation platform. To enter, one descends beneath the pier,
into the underbelly of the structure through a series of connecting
pods. A hard shell holds a series of expandable pods containing
portions of the spa that expand and submerge underwater as the
tide rises. Once inflated, they become stable environments, creating a habitable and desirable space in a previously inaccessible
and dangerous place. It is an escape into an unfamiliar environment of undersea creatures.

View of the spaces beneath the pier. Each pod is inhabitable by one or a few individuals and offers a unique perspective of the environment surrounding the viewer. Depending on the tidal level, the viewer is above water level or
submerged, co-existing with the ocean environment.

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Project Documentation  The project is inspired by the sea star’s
skin system, which is not entirely solid. Between hard and impervious panels, there is a series of soft membranes that pneumatically expand as the tide rises and interfaces with the assemblage.
These membranes inflate and become spaces under water, carving out new real estate and creating a temporarily dynamic interface between humans and an environment typically unsuitable for
them. The walls of this membrane vary in translucency, allowing views of the environment. The temperature of these spaces is
consistently cooler than the external temperatures, ideal for the
peak tourist season during the summer. The membrane is firm
to the touch, yet appears soft. The material composition of the
membrane is a series of layers, the outermost of which is similar
to ETFE. In between there are layers of sponges that when submerged absorb water and displace air.



34 5 67


1. Permeable Membrane
2. Deformable Membrane System
3. Sponge Layer
4. Rubber Membrane




5 67


5. Air Chamber
6. Membrane with Controlled Pores
7. Scented Air Chamber
8. Spongy Bubble Inflatable System

Here a wall section detail depicts the intricacy of the envelope composition. The intent of the architectural envelope
is to provide maximum sensory experience. The soft membrane is composed of multiple layers capable of exchanging volumes of water and gasses with the surroundings, varying the wall thickness when above or below water.

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16 8








The sectional drawing demonstrates how the system hangs, anchored to the underbelly of the pier. It is not unlike
the sea star’s capability to cling to the rocky shores of the coastline. The pillowed spaces are dynamic environments
with no defined floor, no wall, no ceiling, but rather an encompassing padded hollow space which can be utilized
in myriad ways from standing, to sitting, to sprawling. All of these details mimic the experience and function of the
sea star abstracted in the form of a multi-sensory environment.

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As a contrast to the amusement park environment above, the space of the intertidal zone opens up real novelty —
the novelty of experiencing an environment not designed for the user, where architecture protects and allows for
impossible interactions. This becomes a region of water and mist, of dappled light and shadow, alien creatures, and
strange interactions. It is a harsh environment full of perils to its inhabitants. The project creates a calm space within
this tumultuous environment.

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Animal Examples  Gila monsters are large, desert-dwelling lizards that spend almost all of their time in underground burrows,
rarely coming out to feed. These traits allow gila monsters to prevent water loss in the arid habitats in which they live. Horned
lizards have evolved a unique method of collecting water during desert rains. They possess narrow channels in between their
scales that cause water to flow, through capillary action, into their
mouths. Kangaroo rats live in burrows underground that they
cover with dirt during the day to prevent water loss. While they
sleep kangaroo rats bury their noses into their fur to recycle water
as they breathe. Honeypot ants construct nests far underground
where the soil is cool and moist. Some ants in each colony act as
specialized storage devices; their abdomens are able to swell to
the size of marbles in order to store nectar, which provides nutrients and water to other individuals in the colony.

Desert-dwelling animals cope with low availability of water using adaptations that help them acquire water and
prevent them from losing water. (Photographs courtesy of: gila monster, Wikimedia; horned lizard, J. Rothmeyer;
kangaroo rat, S. Yeliseev; honeypot ants, T. Doyle)

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Namib Desert beetles

Onymacris unguicularis, Physasterna cribripes



Genus: Onymacris, Physasterna
O. unguicularis,
P. cribripes
Photograph courtesy of A. Sosio

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Habitat & Climate  The Namib Desert, situated on the western coast of Africa in Angola, Namibia, and South Africa, is
the world’s oldest and one of its harshest deserts. The desert is
broadly characterized into two regions, the coast and inland.
The temperature regime varies between the coast from 9–20ºC
(48–68ºF), and inland from 0–45ºC (32–113ºF). Precipitation in
each region comes from rainfall, but in the coastal region it also
comes in the form of dense, enveloping fog. While the inland
region receives about 5 mm (0.02 in.) of precipitation per year,
the coastal region can get up to 200 mm (7.8 in.) in the wettest
areas. Rain and fog are the only opportunities for desert life to
acquire water since almost no bodies of water are located in the
Namib Desert. The diversity of animal life is composed mainly
of arthropods, notably darkling beetles, commonly referred to as
Namib Desert beetles, adapted to hot and dry desert conditions.


The project is set in the coastal region of the Namib Desert in the country of Namibia. This area is characterized
by vast, undulating sand dunes colored pink and orange. Vegetation is sparse and consists of lichens and succulent
plants near the coast and shrubby vegetation farther inland. (Photographs courtesy of A. Sosio)

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17 3

Animal Physiological, Behavioral & Anatomical Elements 
Due to an extreme scarcity of water, some desert beetles have
evolved strategies to harvest water from the environment. The fog
that arrives in late evening and early dawn represents a more reliable source of water than rain, so beetles have ways to collect
fog droplets. These include sucking water from sand covered in
fog, drinking from water that has collected on vegetation, creating trenches in the sand to collect fog, and finally, fog basking.
Fog basking is a behavioral strategy that involves collecting water
using body surfaces. The beetles face into the wind and assume
a tilted position with their heads close to ground, permitting fog
droplets to accumulate on their backs and trickle down the body
into their mouths. Additionally, they often have long legs to help
with movement across and within sand as well as to raise them
off the hot substrate.
Water droplet accumulates
on tilted body
Exoskeleton ridges
Long appendages elevate
body from ground

Water droplets

Two species of Namib Desert beetles, Onymacris unguicularis and Physasterna cribripes, are nocturnal, burrowing
deep under the sand during the day and emerging at night to feed. Physiological adaptations include excreting very
little water in their waste products and having a low metabolic rate. Many of these beetles also have a waxy surface
over their exoskeletons to prevent the loss of water. (Photograph courtesy of Nørgard and Dacke [2005]; project
team: E. Chen & C. Rodriguez)

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Interface between the Skin & External World  Two species
are known to fog bask: O. unguicularis and O. bicolor. Previous
studies have examined the mechanism by which fog is collected.
There are regions on the elytra surfaces that are coated in wax
(troughs), making them hydrophobic (repellant to water), and
regions that are not covered in wax (peaks), which confer hydrophilic (water-attracting) properties. The small fog droplets congregate on the hydrophilic areas until they form large enough drops,
which funnel down to the mouth. Numerous biomimicry studies
have been based on this water collection mechanism; however,
they were incorrectly based on a species that does not fog bask.
Nevertheless, the physical properties of the hydrophilic peaks that
attract fog droplets have been very successful in designing prototypes and products. This chapter studies O. unguicularis, known
for collecting fog, and P. cribripes for its troughs and peaks.






The surface of P. cribripes with peaks and troughs was used as the inspiration for the project. The bumpy surface of
the elytra is shown with the wax-free, hydrophilic and waxy, hydrophobic areas. Though not biologically accurate,
the physical properties of the water collection mechanism can inspire design. A cross section of the beetle exoskeleton showing the waxy layer. (Photographs courtesy of: bump stain and micrograph, Parker and Lawrence [2001];
dorsal view, Nørgard and Dacke [2005])

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Interrelationship between the Skin & Internal Systems  Insects
breathe using a tracheal system, a series of branching tubes that
connect to the external environment through openings called spiracles. Tracheae branch into smaller tubes called tracheoles that
conduct oxygen directly to every cell in the body. During respiration oxygen is gained but water is also lost through evaporation. In
the desert any lost water comes with a huge cost. Therefore, their
respiratory systems have evolved to prevent water loss. Namib
Desert beetles have lost the ability to fly; their outer wings, elytra,
have been fused together to form a hardened surface with a cavity underneath. The beetles breathe into the cavities, which helps
reduce respiratory water loss. During the day they burrow into
sand to further prevent water loss. Finally, they have long periods
where they stop breathing to minimize the loss of water.






O. unguicularis

P. cribripes

The efficiency in water retention varies among the different beetle species. This project takes two of the most effective beetles and combines the geometry in order to optimize water retention capabilities. The first is O. unguicularis,
in which we mimic the ridges and valleys of the shell, and the second is P. cribripes, which is the inspiration for the
bumpy texture. (Photograph courtesy of Nørgard and Dacke [2005])

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Proto-Architectural Project  The project, located in the Namib
Desert, serves as a temporary residence for desert researchers.
Taking cues from the beetles’ ability to collect water from such
an arid climate, the building is positioned so that the largest
sloped roof/façade is facing southeast toward the morning fog that
sweeps up from the ocean, and burrows a portion of the building
underground to regulate its thermal mass.
The massing takes after a combination of two different species’
exoskeleton structure in order to optimize the water retention
capability. The bumpy texture is inspired by P. cribripes as a
way to capture the moisture in the air, while the overall geometry
was extracted from O. unguicularis with the ridges and valleys to
channel the water into storage.

The project’s geometry allows for a large portion of the façade to face the direction of the morning fog, with an angle
designed so that the water can travel down from the top toward the water storage. Utilizing the characteristics of the
beetles’ bumps and ridges, the highly articulated building’s surface has the capacity to capture water. The water can
then be used to create a microclimate for its inhabitant.

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17 7

Project Documentation  The building envelope uses a series of
mesh discs to capture moisture from the morning fog. Each disc
is set on a pivot and and tilts when enough water has accumulated. The water rolls down the grooved roof membrane toward
the water cistern at the base of the structure. The collected water
not only provides a potable water source for the occupants but is a
cooling mass to help regulate comfort within the space. Not only
does the shape of the building mimic the “fog basking” water collection feature of the beetle, architecturally the form creates an
interior volume (low to high ceilings) which optimizes air stratification and overall climate comfort. A portion of the roof discs
are perforated to allow for the collection of water to sustain the
interior plants. The plants allow for storage of water, food, aromatherapy, and help humidify the space through evapotranspiration.

Large size mesh disc
Medium size mesh disc

Small size mesh disc

High ceiling accumulates and
exhausts hot air

Radiant cooling surface
Rain water storage

The diagrams show the components of the mesh disc and how it collects water droplets. The schematic building
section indicates the exterior positioning and decreasing sizes of the discs as well as delineating the interior space,
slightly sunk into the sand to take advantage of its thermal mass.

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To capture
morning fog




Valleys channel
water drop to irrigate
the vases below and
replenish the
building's cistern
Holds herbs and
plants rooted in
coconut and jute
fibers to generate
a microclimate

Mesh to tilt
from weight
of water

The wall section and rendering demonstrate how water is captured through the discs made out of mesh. The water
is then funneled into the channels by the geometry of the overall massing. The channels are connected to the upside
down pots in the ceiling which grow a range of mixed herbal vegetation. The vegetation shown on the section creates a ceiling condition that is unexpected for a desert condition and provides a small quantity of edible greens.

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In this hot, arid and typically deemed uninhabitable environment, the project provides an unexpected oasis for
researchers. Furthermore, the dynamic system on the roof engages in water collection, transforming the interior with
landscapes of edible vegetation, thus creating an environment that allows the researcher to thrive in such a severe

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Many of the skin functions we have previously considered could
be interpreted as protective adaptations. For example, animals use
communication, often through coloration, to prevent themselves
from getting eaten by a predator or attacked by a competitor.
This may occur in the form of warning coloration or camouflage.
Animals continuously thermoregulate to protect themselves from
environments that are too hot or cold. They also use their skin to
protect themselves from losing water. In this section, we will consider two additional ways in which animals protect themselves —
by using physical defense to protect themselves from attack, and
by chemical defense from predators and the environment.
The ant species Cephalotes atratus
protects and defends its colony by
using heads of the workers to block
its nest entrances. (Photograph courtesy of S. Price)

16_Protection.indd 181

The skin and its appendages provide the first line of defense for
most animals, and many have evolved extreme modifications of
the skin that function in protection against predators or against
members of their own species. These include horns, antlers, sharp
teeth, legs, claws, spines and sharp hairs. The skin itself can be
hardened to serve as protective armor. The shells of turtles, snails,
and clams are examples of hardened animal coverings that prevent predators from getting to the vulnerable, internal body structures. Scales, bony plates and fur are existing appendages born
from the skin that are used to protect from the elements and from
predators. Animals also use behaviors in conjunction with skin
appendages for protection. For example, erection of feathers or
fur by tiny muscles in the skin can make an animal appear larger
than it really is and scare potential predators or competitors.
A vestige of this innate behavior is retained in humans – hairs

13/12/12 11:56 AM


standing up on the back of the neck during a scary movie are
our body’s attempt to appear intimidating to an imaginary threat.
Many animal species move in groups, or herds, as a protective
defense. Their coloration patterns may work in tandem to confuse
Toxins, venom and poison are all chemical defenses that often
are secreted by glands in the skin or protrusions from the skin.
In many animals these secretions are used for scent marking,
but in other cases they are modified to aid in defense from
predators. For example, skunks are able to forcefully eject noxious secretions from their anal glands when threatened. A lot of
amphibian species have poison glands in their skin which, combined with bright warning colorations, deter predators. A lot of
invertebrates secrete chemicals from their joints or specialized
glands that are irritating, toxic, waxy, or odorous. Invertebrates
are often brightly colored to warn predators of their distastefulness. Some marine invertebrates, like octopuses and squid, produce colored pigments that can be ejected to confuse predators
and facilitate their escape.

(Top) Tent-making bats protect
themselves from rain by sleeping
under large leaves that they modify
into a tent-shaped form.
(Bottom) Many lizards, such as the
banded gecko, will undergo the radical process of dropping their tails to
escape predation, leaving the predator with a writhing tail and no prey.
(Photographs courtesy of: bats, S.
McCann; gecko, J. Rothmeyer)

The fundamental idea of shelter is one of a structure that provides
protection from predators and the elements. Over time, the type
and meaning of human shelters have been added to. Shelters were
initially used as protection from the weather, predators and other
human populations. However, shelters not only serve the function of protection, they also serve as a tool of communication. A
primary signal shelters send is that of power. For example, castles
and defensive forts convey power even without actively being
used. Today, structural security intended as defense and safety
occurs in a few specialized building types such as embassies, jails
or courthouses.
Buildings still serve to protect humans from environmental elements,
but now they also need to address more severe challenges such as
ozone layer depletion, air pollution, acid rain, and an increased severity of weather conditions resulting from climate change.
One critical environmental element is the sun, our primary source
of energy. While the sun’s radiation can be turned into an unlimited energy source, it can also produce chemical changes detrimental to humans and the Earth’s atmosphere.

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The trees provide shade and through evapotranspiration mechanisms cool the
protected enviroment. The photovoltaic panels supply the energy necessary
to activate the misting cooling system. Ecosistema Urbano, Ecoboulevard,
Madrid, 2005. (Photograph courtesy of D. Terna)

The problems created by the necessity to both shield and harvest
the sun’s radiation provide unique opportunities in architectural
exploration. From the sun we can acquire energy, and through the
design of a smart active building envelope, mediate it, filter it, and
transform it for internal use and comfort. The use of advanced
materials can further enhance envelope performance. For example, it can include a process involving the sun as a catalyst that
is used to produce coverings that contain titanium dioxide. The
coverings break down inorganic and organic pollutants and also
actively participate in cleaning the air while protecting a building

Three large artificial trees respond to
the scarcity of nature in this peripheral city development, while providing climatic comfort in public spaces.
Ecosistema Urbano, Ecoboulevard,
Madrid, 2005. (Photograph courtesy
of D. Terna)

16_Protection.indd 183

Here we have used two examples to show different ways of taking
insights from nature to design protective structures. Inspired by
the pangolin’s hard scales, the project is a portable structure that
accumulates solar energy to be used in emergencies while protecting inhabitants from the sun’s UV rays. The sweat glands of
hippopotamuses secrete a substance that protects them from UV
rays and helps maintain the animal’s water balance. The designed
building envelope is constituted of tessellated elements that collect water; these are used to create a thermal mass to serve as heat
protection and a water reservoir.

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18 4

Animal Examples — Protection  Animals have a multitude
of mechanisms that serve as protection mainly from predators,
but also from the elements and individuals of their own species.
Meerkats are social animals; when outside of their burrows one
will stand sentry to watch out for predators and alert the other
individuals. Hedgehogs roll into a ball and have stiff, hollow hairs
they use to protect themselves from predators. Skunks are most
known for the noxious liquid they spray from anal scent glands
to ward off predators. Lionfish protect themselves through their
venomous spines. Porcupine fish also have spines; they take in air,
and their bodies inflate, thus protruding the spines. Similarly, sea
urchins have spines that serve as protection. The flamingo tongue
snail is brightly colored due to the live tissue that covers its shell.
When attacked the tissue retreats into the shell.

All animals have adaptations that serve as protection from predators, the environment and members of the same
species. These include spines, toxins, hardened coverings and social behavior. (Photographs courtesy of: meerkats,
I. Mazzoleni; hedgehog, Wikimedia, J. Hempel; skunk, B. Garrett; lionfish, A. Jaffe; porcupine fish & pencil urchin,
J. Maragos; flamingo tongue snai, A. Jaffe)

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Animal Examples — Protection  Spiders, such as black widows
and orb weavers, have venom to subdue prey but also to protect
from predators. The armadillo lizard will roll into a ball, hence
the common name, by putting its tail in its mouth and protecting
itself with thick, sharp scales.
Harpy eagles are top predators; they use their sharp claws to capture prey. Their coloration patterns help them blend into the forest
canopy to sneak up on their prey. Pill millipedes roll into a ball
when disturbed. Rhinoceros beetles are large beetles with very
thick exoskeletons. Males use horns in combat with other males.
Many animals, such as butterflies, have eyespots, markings on
their bodies that resemble eyes. When flashed they are thought to
mimic other animals to deceive predators.

All animals have adaptations that serve as protection from predators, the environment and members of the same species. These include spines, toxins, hardened coverings and social behavior. (Photographs courtesy of: black widow
spider & orb weaver spider, M. Hedin; armadillo lizard, P. le FN Mouton; harpy eagle, A. Kirschel; pill millipede, M.
Hedin; rhinoceros beetle, S. Yanoviak; butterfly, S. McCann)

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18 6

Animal Examples  Mammals have evolved many ways of protecting themselves, both from members of their own species and
from potential predators. Wildebeest live in large herds and use
speed as a primary means of defense. They also possess horns,
which can be used as a last line of defense from predators, as well
as for fighting members of their own species for access to mates.
Other animals use coverings as protection. Porcupines are covered in quills, which are actually keratinized hairs and deter
predators. Armadillos are protected by a series of bony plates
covering the back, head and tail. When attacked, armadillos roll
into a tight ball, protecting the vulnerable belly. Lions may not
seem as though they need protection, but the shaggy mane covering the neck of the male may act as protection when they engage
in to-the-death fights over access to females.

Mammals use behaviors to protect themselves as well as modifications to their skin and protrusions from their skin.
(Photographs courtesy of: wildebeest, Wikimedia, C. Rosenthal; porcupine, L. Parenteau; armadillo, A. Patterson,; lion, B. Larison)

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Tree pangolin

Manis tricuspis


Chordata Family:
Mammalia Genus:
Pholidota Species:

M. tricuspis

Photograph courtesy of E. Simpson

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18 8

Habitat & Climate  There are eight species of living pangolins,
all within the genus Manis. The tree pangolin, a small African
species, inhabits areas with suitable tree cover such as woody
savannahs and the lowland tropical rainforests of western Africa.
The location for the project is the Kakum National Park in Ghana,
located in the heart of West Africa and home to lush tropical rainforest in less than 350 km2 (144.8 mi2), containing many of the
world’s most endangered plant and animal species. The average
yearly air temperature ranges from 26 to 33°C (~79–91°F); the
park receives a high amount of precipitation, an average yearly
amount of 1100 mm/yr (43.3 in.), corresponding to ~57 days of
rain. The park’s morning humidity averages 96% while the evening average is 79%. The average hourly direct normal radiation
range is 425 to 75 Wh/m2, and the average hourly direct normal
illumination range is 35,000 to 5000 lux.


Habitat images of Kakum National Park, Ghana, West Africa. Tree pangolin distribution map. (Photographs courtesy
of I. Mazzoleni)

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Animal Physiological, Behavioral & Anatomical Element 
Tree pangolins weigh 4.5–14 kg (10–31 lbs) and have body lengths
of 31–45 cm (12–18 in.). The pangolin’s sides and back are covered with large, overlapping scales composed of keratin, similar
to human nails. When threatened, pangolins curl into a tight ball,
so the scales provide a layer of defense from predators and stinging insects, as they feed exclusively on ants and termites. Due to
their lack of teeth, they possess tubular snouts and long tongues.
At rest, the tongue is drawn into the throat by long powerful
muscles originating from the sternum. Unusually large mucusproducing salivary glands lubricate the tongue and provide an
adhesive surface on which ants can be trapped. Tree pangolins
spend much of their time in the trees and have developed a large,
muscular tail that serves both as a counterbalance and a fifth limb
to aid in climbing and hanging from branches.





Inflexible, hard

Long, sticky
Flexible, muscular and defensive

The main morphological characteristics of the tree pangolin include protective scales, large retractable claws for
climbing and opening termite nests, and a long, sticky tongue which allows the pangolin to reach deep into termite
and ant mounds. The scales are grooved or corrugated on their surface. In burrowing pangolins, this protects against
abrasive damage from dirt and rocks. In tree pangolins, it may function to protect from scratching branches and
thorns. (Photograph courtesy of K. Benirschke; project team: R. Ferrari & T. Carpentier)

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19 0

Interface between the Skin & External World  Like all mammals, the pangolin’s skin is composed of three layers: the hypodermis, dermis, and epidermis. However, the overall physical
appearance of the pangolin is dominated by large, hardened, platelike scales that protect the animal from predators and prey. The
scales are actually keratinized cells that grow from raised papillae
protruding from the surface of the skin. The scales of newborn pangolins are soft but harden as they mature. The exposed outer and
inner surfaces of the scales may fray through wear, but are replaced
by newly keratinized cells from the middle layer.
Critically, the pangolin’s skin system provides a hard protective
system that maintains sufficient flexibility for the animal to climb
or dig. The scales have a corrugated surface, which prevents
excessive localized damage from everyday wear and tear.





The illustration shows a detail of a cluster of the pangolin’s scales. The skin and scale section show the connection of the scale to the skin and the raised papillae from which the scale growth occurs. (Photograph courtesy of K.
Benirschke; diagram adapted from D. Visset; copyright credit Editiones Belin, 2006)

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Interrelationship between the Skin & Internal Systems  The
pangolin is often compared to a walking pine cone or globe artichoke. Although they tend to move at a slow pace, the flexibility
afforded by scale armor, as opposed to a solid shell, means that
the pangolin can move much faster when needed. When threatened, they are able to rear up on their hind limbs while utilizing
the tail for balance. This increases the size of the pangolin and
exposes the large claws in a warning threat display.
When threatened and unable to defend itself, the pangolin will roll
up into a ball, using its prehensile tail to cover its face. Pangolins
can also produce a noxious secretion from anal glands, which
may also scare off a predator. In conjunction with protective eyelids, the pangolin is able to resist the attacks of soldier ants and
termites and feed with relative ease.

Walking position

Defense position

The diagram illustrates the pangolin’s retractable claws, which allow them to walk on their front knuckles with the
claws tucked underneath to protect them from wear, similar to giant anteaters. The claws also provide them with
an additional protective element. The pangolin’s tail plays a crucial role in movement and protection. (Photographs
courtesy of: climbing, A. Kirschel; hanging, D. Ellis)

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Proto-Architectural Project  This project explores the duality
of the protective and flexible qualities of the pangolin skin system, juxtaposing a flexible skin made up of rigid scales. Using
Kakum National Park in Ghana as an initial project location, the
building program evolves into an emergency pavilion for postnatural disaster or war zones. The project focuses on a premise to
construct and develop a deployable and temporary structure that
would adapt to varying site conditions and terrains. In the given
scenario the primary interest is to establish temporary shelters
to assist medical teams. The panels are held together using the
principle of tensegrity, to provide the structural capacity necessary to support the panels and produce an inhabitable environment. In this instance the panels are lifted and maintained in their
positions through tension and compression applied through the
structural posts and cables.

An initial study model explored the flexible geometry of origami paper folding, especially with regard to the notion
of expandable geometry. The examination led to the development of a final study model of a shelter which accommodates people and different programmatic activities. The final shelter design may be expanded and contracted on
site to respond to different conditions such as light porosity, area, volume, and whatever degree of environmental
protection is necessary.

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Project Documentation  The shelter is designed for transportability in a series of basic pieces: photovoltaic panels, flexible translucent fabric, and a connecting umbrella-like structure.
Whereas the pangolin’s skin system provides protection against
prey and predators, the adaptability of this shelter focuses on providing protection from undesirable environmental conditions. To
address the necessity of energy, the design clads the modules with
photovoltaic panels so that energy becomes readily available.
Assembly of the shelter is a multi-step process that begins by
expanding the structural umbrella to stretch the protective fabric. Next, the photovoltaic panels are attached to the basic unit
modules, which are then attached to form a stable arch, creating
structural rigidity for the shelter. Finally, rows of unit modules are
repeated until the desired volumetric space is achieved.

ground connection

maximum solar yield diagram

photovoltaic panel
flexible fabric


A series of assembled modules lock in place to form a stable arch. The rigid photovoltaic panels are oriented to
act as a protective shading element as well as to capture solar energy for use in the shelter. Right middle: singular
assembled module. Right bottom: exploded diagram of an assembled module.

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19 4





The diagrams illustrate the flexibility derived through the shelter’s kit-of-parts assembly. En route to a disaster, the
shelter elements are disassembled and shipped in small containers to make transport more effective. By opening or
closing the assembled modules, the shelter envelope can be expanded or contracted to adapt to surface area and
volume needs.

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Inspired by the protective and flexible qualities of the pangolin skin system, the nomadic modular shelter provides
protection from environmental elements while producing electricity for its inhabitants. The translucent envelope
made of fabric allows an abundance of light and natural ventilation to pierce through, while maintaining a rigid
tactility through principles of tensegrity.

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19 6

Animal Examples  Mammals, including humans, keep cool in
a number of ways. Our skin is hairless, allowing air to flow freely
over it. Sweating causes body heat to be lost via evaporative cooling. Tanning results from increased melatonin production to protect from UV radiation. Warthogs are also mostly hairless. They
spend the hottest parts of the day in cool underground shelters
and may wallow in mud to provide additional cooling. Spotted
hyenas are covered in fur. Their fur is pale in color to reflect UV
radiation and minimize overheating.
When hyenas do overheat, they seek shade and pant, like domestic dogs, to lose heat via evaporative cooling from the tongue.
Rhinoceroses possess thick, dark, hairless skin. Water loss and
UV damage are minimized by the densely keratinized epidermis.
Rhinos, too, may wallow to further cool themselves.

Animals have to protect themselves from environmental conditions, such as heat. They do this through sweating,
panting, and spending time in the shade and other cool places. (Photographs courtesy of: human skin, S. Yelisee;
warthog, spotted hyenas and rhinoceros, B. Larison)

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Hippopotamus amphibius


Chordata Family:
Mammalia Genus:
Artiodactyla Species:

H. amphibius

Photograph courtesy of B. Larison

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Habitat & Climate  There are two species in the Hippopotamidae
family. The largest species, the common hippo, occurs only in
grassland areas of subsaharan Africa where still water can be
found. Hippos occupy an open stretch of water where they can rest
during the day. At night, they leave the water to graze on grasses.
The Okavango Delta of Botswana is an optimal common hippo
habitat. The Delta is an oasis in an otherwise arid landscape with
a predominantly subtropical climate. Average rainfall is 450 mm
(17.7 in.), most of which falls between December and March during heavy thunderstorms. Seasons vary from hot, wet summers
during December through February to cold, dry winters in June
through August. In the summer, temperatures reach as high as
40°C (104°F) with humidity levels between 50 and 80%. Night
temperatures during winter may reach barely above freezing.

The Okavango Delta


Botswana is dominated by the Kalahari Desert, covering 70% of its surface. The Okavango Delta in the Northwest
offers a reprieve from the otherwise parched environment. During the Delta’s annual flood, the water-covered area
increases almost three-fold, from about 5000 km2 (1930 mi2) to 6000–12,000 km2 (2316 mi2–4634 mi2) during the
winter months. (Photographs courtesy of B. Larison)

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Animal Physiological, Behavioral & Anatomical Elements 
The common hippo is characterized by a barrel-shaped torso,
enormous mouth, large teeth, and relatively short legs ending in
feet with webbed toes that make it easier to move underwater.
The hippo’s skin is almost hairless, with just a few bristles around
the mouth and the tip of the tail, and is especially thick over the
back and rump. Skin color varies from grayish-brown on the back
to pink on the belly and around the eyes and the ears where the
skin is thinnest. The hippo’s thick skin is further protected by
secreting a thick fluid from glands in the skin that protects against
sunburn and keeps the coarse skin moist when the hippo is out of
the water. Hippos also stay under water as a means of protecting
the skin from sunburn and to keep cool. During hot days hippos
come out of the water to graze only at night.

Thick skin over back
and rump

barrel-shaped torso

Eyes and ears on
top of skull

AM - cold

- hot

PM - cold

Most of body submerged
Eyes and ears above water

Front legs

Hind legs

Short legs with
webbed toes

Thinner skin on
belly and limbs

Large muzzle with
nostrils that close
under water
Webbed toes

The eyes, ears and nostrils are placed high on the skull so that they do not submerge. The hippo’s large muzzle has
nostrils that close when underwater, allowing the hippo to fully submerge for several minutes at a time. The large
body mass of the hippo allows it to walk underwater. It cannot float or swim. They move by bouncing off the riverbed
floor and walking on the bottom. Webbing between the toes helps to propel them through the water. (Project team:
S. Månsson & W. Raksaphon)

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20 0





Constant Moisture Level


Thick Dermis



50 µm

The thick skin consists of a dense sheet of collagen with fibers arranged in a matted, but regular, pattern giving great
strength to the skin. Hippos have neither sweat nor sebaceous glands, but they have glands that produce a viscous
red fluid, leading to the myth that hippos “sweat blood.” Hippos secrete a natural sunscreen from glands deep within
the dermis. This substance protects the thin epidermis from harmful UV rays, while also providing some antibacterial
protection. (Micrograph courtesy of Reed et al. [2009]; diagram adapted from Luck and Wright [1963])

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Interrelationship between the Skin & Internal Systems  The
principal physiological functions of the hippo’s skin are to control body temperature and regulate water loss. The hippo’s semiaquatic lifestyle allows it to control both simultaneously. By
remaining submerged in water during the hottest parts of the day,
hippos avoid body warming due to environmental factors. Hippos
also have relatively dense bone, particularly in their limbs, to neutralize the natural buoyancy of the body and reduce the amount
of effort and energy required to remain submerged. During the
dry season when water levels are low, temperature regulation can
occur via the activation of thermal windows. Thermal windows
are areas of the body with momentarily higher temperatures than
the surrounding body surface and ambient temperature. These
windows of warmed skin lose heat to the environment, cooling
the body in the process.

Ambient temp. 21oC

16.2 C

Ambient temp. 28oC

20.8 C

Thermal windows

skeletal structure

Specific gravity allows
the hippo to sink and
run under water
Heart rate slows
down to stay under
water longer

Hippos submerge with their ears pressed down flat and nostrils clamped shut. While underwater, their heart rate
slows so they can stay underwater longer. As hippos feed on land, temperature regulation can be aided by flushing body heat to thermal windows in the skin. Thermal windows are increased in higher ambient temperatures.
(Diagram adapted from Schneider and Kolter [2009])

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Proto-Architectural Project  The project entitled “The Porous
Skin” controls the intake of daylight while managing temperature
during the different seasons. Inspiration for the design is found
from the hippo’s semi-aquatic lifestyle and the protective quality observed in its skin. The project’s building envelope creates
a comfortable indoor climate in both the warm summer and the
colder winter months. The sliding panels regulate both the amount
of daylight coming into the building and the water retained in the
envelope assembly, which acts as a cooling element during the
summer and a thermal mass in the winter. The permeable enclosure varies from a thin almost perforated envelope to a thicker
dense wall, allowing for the transfer of heat from one layer of the
façade to another. The various orientations of the opening within
the wall unit control the direction of the light seeping into the
building and enable control over the heat intake.

Hippo skin produces protective fluid in response to changing environmental conditions. The panelized façade system is created to accommodate the changing climate of the Okavango Delta which is characterized by warm and wet
summer months and a cold and dry winter season. This design makes it possible for the building envelope to change
through different conditions, always taking full advantage of the varying climate of the building site.

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Project Documentation  The building envelope is composed of
movable panels shifting from one position to another. This creates
a skin system that transitions from an entirely closed envelope
that lets in very little daylight to an open envelope that allows the
sun to heat up the interior of the building. This design accommodates the changing climate of the Okavango Delta.
When the panels are set to position A, water flows into the outer
layer of the skin to cool the interior of the building during the hot
and wet summer season. Opening the structure in the winter season with panels set to position B allows for more sunlight to permeate the façade, and helps warm the living spaces. Waiting until
the peak of winter to set panels to an open position helps retain
water between the layers of the envelope, functioning as a thermal
mass which slowly releases stored heat within the building.
High sun angle primarily in the summer

Low sun angle primarily in the winter
Open segment - Position B
Movable element

Angled façade to control
intake of light

Closed segment - Position A

The intent of the building envelope is to create a comfortable indoor climate in both the warm summer and the
colder winter months. Sensors manage the sliding of the wall units controlling the overall amount of light directly
hitting the floor plate, therefore regulating the temperature in specific areas of the building.

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20 4




of water

Panel - Position A

of light

Panel - Position B

The aluminum panels consist of a base unit with multiple openings. One half of the panels contain vertical apertures
which let in large amounts of sunlight. The other half of the panels has fewer, slightly tilted openings, angled to capture rainwater. When panels slide into the closed position the angled apertures open and become permeable to both
light and water, while the vertical openings remain as apertures, permeable only to light.

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20 5

The panelized façade system creates a dense and layered, functional and ornamental building envelope. Filtering
light through a permeable membrane drastically alters the ambience of the internal spaces as light glimmers among
the internal surfaces fluctuating internal temperatures to comfortable levels. The porous and formed expression of
the façade promotes a unique condition of shifting panels to establish an operable building that opens and closes as
it registers the changes in the surrounding environment.

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1. Biomimicry is a term coined by Janine M. Benyus in her book
Biomimicry. Innovation Inspired by Nature (1997).
2. Oxford Dictionary: Prefixed to nouns and adjectives with the
sense “earliest, original; at an early stage of development, primitive; incipient, potential.”
3. Philip Drew Touch This Earth Lightly: Glenn Murcutt in His
Own Words” (1999).
4. These data do not include species that are extinct in the wild.
For further data on extinction rates see
5. Ibid.
6. SCI-Arc Lecture, March 8, 2012.
7. Ibid.
8. Tom Wiscombe, “Beyond assemblies: System convergence and
multi-materiality,” Bioinspir. Biomim. 7 (2012), IOP Publishing.
9. “Materiomics is an emerging field of science that provides a
basis for multiscale material system characterization, inspired

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20 8

in part by natural, for example, protein-based materials.” From
Materiomics: Biological Protein Materials, from Nano to Macro
S. Cranford and M.J. Buehler, Dovepress (2010).
[10] In humans, this organ serves as a protective barrier against
microorganisms, physical damage, UV radiation, water loss, and
the elements. It helps control body temperature through evaporative cooling, heat retention, radiation, convection, and conduction.
It aids in fighting infections by providing an acidic environment
and anti-microbial secretions and through initiating the body’s
immune response and the healing process.
[11] Many animal species are sexually dimorphic, meaning that
males and females take different forms. Often, drab-colored
females will be attracted to males that are vividly colored, because
bright colors are thought to be an indicator of male health. An
evolutionary advantage may be gained by females who mate with
healthier males, as their offspring are likely to be healthier.
[12] In vertebrates, pigments are found in specialized branching cells called chromatophores. The distribution of the pigment
within the cell is generally what causes dark coloration (when pigments are distributed throughout the cell) or light coloration (when
they are concentrated in one portion of the cell). Coloration also
has to do with the amount of pigment and the rate it is produced,
or a change in the shape of the chromatophores. Invertebrates
share the same pigments as vertebrates, but they are not always
contained within specialized cells. Brown or black colors result
from the presence of melanin; its stable chemical structure helps
protect the skin from abrasion, and it absorbs UV light, which can
damage tissue. Carotenoid pigments on their own provide yellow,
orange, and red coloration; however, they are sometimes associated with different proteins, which can cause almost any kind of
coloration. Carotenoids cannot be manufactured by animals, so
they need to be obtained from plants. Finally, pterins are colorless, red, or yellow pigments that occur mostly in insects. When
they do occur in vertebrates, pterins are often associated with
melanin or carotenoids and contribute to yellow, orange, and red

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[13] Some contain chemicals, such as guanine, that contribute
to white coloration. Other chemicals reflect light and can work
with chromatophores to produce iridescent colors. Animals that
can rapidly change their coloration, such as fish, amphibians, and
reptiles, are usually able to do so by regulating the concentration
of pigments within chromatophores.
[14] Layers of air pockets or of certain kinds of particles in the
skin occur in layers, and the interference between light waves is
reflected in different layers. Interference occurs when light waves
reflect off surfaces and coincide with each other, leading to a perception of brilliant color. The thickness and spacing of the layers
determine which color will be seen. Iridescent colors are intense
but change with the angle of view and can disappear entirely
when viewed from a different direction. The most common iridescent colors are blue and green, but they can also be violet,
gold, orange, and red. Often, pigment and light-scattering layers
are combined to produce exceptionally pure colors. White is usually a structural color where all wavelengths of light are reflected
from the surface.
[15] Additionally, birds use an oil secreted from the uropygial
gland to preen feathers, a process that aids feather interlocking.
[16] A high salt concentration in the environment can cause water
to be lost and desiccation to occur. Freshwater has a lower salt
level than body tissues, causing water to seep into cells, ultimately
causing them to burst. Aquatic organisms and even amphibians
have various physiological mechanisms to maintain this balance.

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Author Biographies

(Photograph courtesy of E. Oprandi
& S. Yanoviak, respectively)

Ilaria Mazzoleni is an architect and the founder of IM Studio
Milano/Los Angeles. Her conceptual work has been published
globally, and her built work can be found in Italy, California,
and Ghana. Ilaria has gained attention in the fields of sustainable architecture and biomimicry. This has led to her being
invited to participate in multiple international conferences and
workshops and her written contributions are published in several
international architectural magazines. Since 2005 she has been
a full-time faculty member at the Southern California Institute
of Architecture (SCI-Arc) in Los Angeles. Her professional and
academic investigation relates to sustainable architecture on all
scales of design with a research focus on biomimicry, where
innovation in architecture and design is inspired by the processes
and functions of nature. Collaborating with biologists and other
scientists from top research institutions, her projects explore the
connections between biotic and abiotic elements within ecosystems in order to develop sustainable urban planning strategies and address solutions to global climate change. An ongoing
research program has centered on understanding how organisms
have evolved and adapted to their environment, and applying that
knowledge to design building façades. Current investigations
use design as a vehicle to promote awareness about endangered
species and emphasize the importance of biodiversity in regions
around the world. Mazzoleni explores the performative capacities
of organic systems using an analytical approach and the strategy
of juxtaposition of real and digital space. The conceptual implications arising from biomimetics and design have led to a body
of work that investigates innovative material processes, forms,
geometries and structural patterns.
Shauna Price is an evolutionary biologist focusing on speciation
in neotropical insects. Her research examines the historical and

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2 30

ecological factors contributing to the high species diversity found
in ants with the use of genetic tools, geological data, and morphological analyses. In addition to conducting research, Shauna
has collaborated with Ilaria Mazzoleni and IM Studio MI/LA on
multiple bio-inspired design projects. She contributes a strong
background in ecology and evolution to these studies, with the
perspective that inspiration in architecture and design can stem
from organisms as small as microbes to broad, ecosystem scales.
In particular, symbiotic relationships—close, ongoing associations that have co-evolved between different species—inform her
perspective in merging biology with design.

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Skin is a complex organ that performs a multitude of functions; namely, it serves
as a link between the body and the environment. Similarly, building envelopes
act as interfaces between their inhabitants and external elements. The resulting
architectural designs illustrate an integrative methodology that allows architecture
to follow nature.

“Ilaria Mazzoleni, in collaboration with biologist Shauna Price, has developed a profound
methodology for architectural and design incentives that anticipates and proposes novel
ways to explore undiscovered biological inspirations for various audiences.”

-Yoseph Bar-Cohen

Architecture Follows Nature

Biology influences design projects in many ways; the related discipline is known as
biomimetics or biomimicry. Using the animal kingdom as a source of inspiration,
Ilaria Mazzoleni seeks to instill a shift in thinking about the application of biological
principles to design and architecture. She focuses on the analysis of how organisms
have adapted to different environments and translates the learned principles into
the built environment. To illustrate the methodology, Mazzoleni draws inspiration
from the diversity of animal coverings, referred to broadly as skin, and applies them
to the design of building envelopes through a series of twelve case studies.


Applying Properties of Animals Skins to Inspire Architectural Envelopes

Ilaria Mazzoleni

in collaboration with Shauna Price

Architecture Follows Nature

ISBN: 978-1-4665-0607-7


9 781466 506077

K14605_COVER_final.indd 1

12/7/12 11:06 AM


What is biomimicry design in architecture? ›

Biomimicry in architecture and manufacturing means designing buildings and products to mimic or co-opt naturally occurring processes. Evolution has shown how organisms have adapted to specific environments, exhibiting resource management that can be a lesson to designers.

Why is biomimetic architecture important? ›

Utilizing the biomimetic principles in architecture design leads to the development of the required and attractive characteristics of the building product such as adaptive architectural envelopes, optimum lighting to spaces, healthy inspired environment, beautiful, sustainable and green surroundings [6].

What is a biomimetic design? ›

Biomorphism refers to designs that visually resemble elements from life (they “look like” nature), whereas biomimetic designs focus on function (they “work like” nature). Biomorphic designs can be very beautiful and beneficial, in part because humans have a natural affinity for nature and natural forms.

What is nature inspired architecture called? ›

Biomimetic Architecture

The design of a building must be carefully considered to improve its functionality and its environmental impact. Biomimetic architecture is the process of imitating the aspects of nature that work well and applying them to elements of the design and construction of buildings.

What are the 3 types of biomimicry? ›

"There are three types of biomimicry - one is copying form and shape, another is copying a process, like photosynthesis in a leaf, and the third is mimicking at an ecosystem level - like building a nature-inspired city."

How can we use biomimetics in architecture? ›

Biomimicry in architecture is often used to seek sustainable measures by understanding the principles governing the form rather than replicating the mere form itself. It applies to several aspects of the architectural and engineering field in terms of materials, structural systems, design, and much more.

Is biomimetic architecture sustainable? ›

Biomimetic architecture is a multi-disciplinary scientific approach to sustainable design that goes beyond using nature as inspiration for aesthetics but rather deeply studying and applying construction principles that are found in natural environments and species.

How are biomimetic materials made? ›

Biomimetic materials have been developed by conjugating synthetic peptide sequences to biomaterials. Growth factor-derived peptides can be directly applied to the polymeric biomaterials.

What is meant by biomimetic? ›

: the study of the formation, structure, or function of biologically produced substances and materials (such as enzymes or silk) and biological mechanisms and processes (such as protein synthesis or photosynthesis) especially for the purpose of synthesizing similar products by artificial mechanisms which mimic natural ...

What are biomimetics used for? ›

Biomimetics is the study of nature and natural phenomena to understand the principles of underlying mechanisms, to obtain ideas from nature, and to apply concepts that may benefit science, engineering, and medicine.

Why is biomimicry so important? ›

The application of biomimicry can benefit the built environment through site design, construction, and operations, as well as reduce the negative impact on the natural environment of numerous techniques for reducing carbon emissions, waste, and others.

Why is nature important in architecture? ›

Natural light plays on the comfort, health, and mood of the human. The design of a building is an integral part of architecture as it brings added value. Making nature visible within a building elevates the spirit. It serves as a visual connection between the indoor, the outdoors, and the natural environment.

What kind of design is inspired by nature? ›

This approach to human innovation, via emulating nature, is called biomimetic design and has inspired many of our greatest creations - from buildings to bionic cars, here are some of the favourite examples.

What are the influences of nature that affects architectural design? ›

The natural surroundings of a particular region influence the colours, materials, spatial planning, structural systems, etc., of a structure. Including architecture interiors according to the regions will allow it to blend in with the surrounding and also help deal with factors like climate and other site features.

What are the principles of biomimicry? ›

The key idea of biomimicry is to learn from nature to solve human problems: learning how to solve human problems, how to invent new things (or at least think we need) and how to live sustainably.

What are biomimetic HOW IT design and which are the most relevant examples? ›

Biomimicry is the science of applying nature-inspired designs in human engineering and invention to solve human problems. It was used to create the first flying machine, inspired by eagles and owls—this paved the way for technologies like jets and planes.

What is interesting about biomimicry? ›

Biomimicry can help emulate and enhance the ecosystem. Designing urban environments that imitate the services of a natural ecosystem like rainwater harvesting, carbon sequestration, energy production, etc. can create an environment that compliments and contributes to the ecosystem.

When did biomimicry in architecture start? ›

1997: With her groundbreaking book, Biomimicry: Innovation Inspired by Nature, Jenine Benyus coined the term biomimicry and sparked the interest of the subject into engineers and designers all over the world. She also started her own world leading consulting organization, Biomimicry 3.8.

What is Biophilic design in architecture? ›

Biophilic design is an approach to architecture that seeks to connect building occupants more closely to nature. Biophilic designed buildings incorporate things like natural lighting and ventilation, natural landscape features and other elements for creating a more productive and healthy built environment for people.

Why is it important for architects and engineers to consider the gift of nature in their design? ›

Why is nature important in architecture? A. Making nature visible within a building elevates the spirit. It serves as a visual connection between the indoor, the outdoors, and the natural environment.

What are ways architects imitate nature? ›

Dear Akram, architects can imitate nature through biomimicry, solving problems by emulating nature through observing and studying its workings. Some examples of biomimicry: bullet trains inspired by Kingfisher birds, helicopter design by the dragonfly and the humpback whales, Airbus design by the albatross.

What is biomimicry in interior design? ›

Biomimetic design is the process of creating innovative ideas inspired by nature. This approach adapts processes of natural organisms to solve design problems and guides design in interior architecture, similar to many other disciplines.

What is organic form in architecture? ›

Organic architecture is a type of architectural design wherein buildings are inspired by, built around, and blend in with their natural surroundings.

How can biomimicry help in developing new materials? ›

Biomimetic materials research creates numerous opportunities for devising new strategies to create multifunctional materials by hierarchical assembly, for the clever use of interfaces and the development of active or self-healing materials.

Why is biomimicry important in tissue engineering? ›

Biomaterials play a pivotal role as scaffolds to provide three-dimensional templates and synthetic extracellular-matrix environments for tissue regeneration.

What type of material can be designed to mimic the properties of other materials? ›

1.5 Ceramics as biomimetic materials. The biomimetic material is a synthetic material, which resembles or imitate material available in nature and retains its functionality.

What are biomimetic reagents? ›

In relation to the second definition, synthetic organic or inorganic catalysts applied to accomplish a chemical transformation accomplished in nature by a biocatalyst (e.g., a purely proteinaceous catalyst, a metal or other cofactor bound to an enzyme, or a ribozyme) can be said to be accomplishing a biomimetic ...

What is the other term for biomimetic? ›

Biomimetic synonyms

In this page you can discover 8 synonyms, antonyms, idiomatic expressions, and related words for biomimetic, like: peptide-based, self-assembling, nanostructures, supramolecular, nano-scale, microfluidics, biosensor and nanostructured.

What is the difference between biomimicry and biomimetics? ›

Bionics: Bionics is the development of a modern system or set of functions based on a similar system that exists in nature. Biomimetics: Biomimetics is the process of mimicking the formation, structure or function of a biologically produced substance or material in order to produce or synthesize an artificial product.

What is an example of a nature inspired technology? ›

Probably the most obvious example of nature-inspired technology is the airplane. It's hard to look at a majestic bird flying through the sky, and not envy its freedom. Humans have been doing it for centuries. So it is not surprising that it's been a goal of humans to learn how to fly for just as long.

How is biomimicry used today? ›

Modern turbine blades form a prominent example of biomimicry in real life. The concept of modern turbine blades is inspired by the structure of the flippers of humpback whales.

How can biomimicry help the environment? ›

Integrating biomimicry into your design practice also can generate multiple benefits for the community at large. Buildings, streets and parks can be constructed to perform the same functions a natural ecosystem does: stormwater harvest; flood mitigation; habitat creation; energy production; and carbon sequestration.

What are the advantages of adopting biomimicry? ›

Creation of green market and services, protection of biodiversity, and conservation of natural resources are the top three benefits established.

What does nature mean in architecture? ›

Nature being as the inherent force which directs either the world or human beings or both and the material world itself, the architects sometimes tend to behave as they are the inherent force and can create anything which can be propagated for their self benefit as part of nature.

What is the nature of architecture? ›

Based on documentation originating in the environmental sciences, history of science, philosophy and art, Architecture of Nature explores the materiality and the effects of the forces at play in the history of the earth through the architect's modes of seeing and techniques of representation.

What is the relationship between architecture and nature? ›

Architects need to make buildings that are friendly to the environment and more green which can be adaptable to the surroundings, in other words, they need to create buildings that are energy efficient, like green buildings or sustainable buildings which are designed to reduce the overall impact of the built ...

What are the natural elements of design? ›

The basic considerations of natural design can be broken down into three categories: aesthetic, managerial, and environmental. The aesthetic aspect of our designs is highly subjective, and individual style varies greatly.

What is nature of design? ›

Design is a systematic action by which solution to the needs of humankind are obtained and communicated. Design is essence of Engineering. Designing is a multidisciplinary task influenced by technological and social factors. Designing is iterative, team work and continually learning process.

What are the key factors in architectural design? ›

Top 5 Factors That Influence Architectural Design
  • Geography, Climate, and Commercial Stair Design. Geography plays a vital role in architecture. ...
  • Religion, Technology, and Culture. Many clients don't have any specific requirements prevailing to religion. ...
  • Imagination and Style. ...
  • Budget. ...
  • Design Changes.
20 Aug 2021

What are the three important influences that we need to consider prior to the design of a building? ›

Context typically encompasses three areas of consideration: physical, cultural, and intended use.

Why is it important to consider environmental factors in our building design? ›

Most of the environmental influences on buildings are variable by their very nature. This fact creates a need for diverse solutions in energy and environmental design and construction that can facilitate optimum building performance throughout the changing periods.

What are some of the best examples of biomimicry in architecture? ›

Here is some example which shows an exemplary use of biomimicry to create some truly unconventional structures :
  • Lotus temple.
  • Lavasa.
  • Hive.
  • Aakash skyscraper.
  • Morarjee Textiles factory.
  • The Tote, Mumbai.

What is the aim of biomorphic architecture? ›

It is a modern architectural style that adopts the idea of embracing natural shapes and patterns into the architecture. Biomorphism aims at turning naturally organic shapes into functional structures.

What is bioclimatic design? ›

Bioclimatic architecture is a way of designing buildings based on the local climate, with the aim of ensuring thermal comfort using environmental resources. They must also blend into their natural surroundings.

What are biomimetic HOW IT design and which are the most relevant examples? ›

Biomimicry is the science of applying nature-inspired designs in human engineering and invention to solve human problems. It was used to create the first flying machine, inspired by eagles and owls—this paved the way for technologies like jets and planes.

Why is biomimicry so important? ›

The application of biomimicry can benefit the built environment through site design, construction, and operations, as well as reduce the negative impact on the natural environment of numerous techniques for reducing carbon emissions, waste, and others.

What are nature-inspired designs? ›

Also known as Biomimetics, it is the interdisciplinary field of creating products by reverse engineering nature. Simply put, it helps researchers study natural phenomena to obtain ideas from nature and apply them to solve real world human problems.

What are the principles of biomimicry? ›

The key idea of biomimicry is to learn from nature to solve human problems: learning how to solve human problems, how to invent new things (or at least think we need) and how to live sustainably.

What is meant by biomimetic? ›

: the study of the formation, structure, or function of biologically produced substances and materials (such as enzymes or silk) and biological mechanisms and processes (such as protein synthesis or photosynthesis) especially for the purpose of synthesizing similar products by artificial mechanisms which mimic natural ...

How do people use biomimicry? ›

Biomimicry, as it's called, is a method for creating solutions to human challenges by emulating designs and ideas found in nature. It's used everywhere: buildings, vehicles, and even materials — so we thought it'd be fun to round up a few of the most noteworthy examples.

What is Biophilic design in architecture? ›

Biophilic design is an approach to architecture that seeks to connect building occupants more closely to nature. Biophilic designed buildings incorporate things like natural lighting and ventilation, natural landscape features and other elements for creating a more productive and healthy built environment for people.

What is a biomorphic shape? ›

The term biomorphic means: life-form (bio=life and morph= form). Biomorphic shapes are often rounded and irregular, unlike most geometric shapes. An artist that loved to explore the possibilities of mixing geometric and biomorphic shapes was Henri Matisse.

What is organic form in architecture? ›

Organic architecture is a type of architectural design wherein buildings are inspired by, built around, and blend in with their natural surroundings.

What are the principles of bioclimatic architecture? ›

[18] described the bioclimatic principles as building design, building envelope, shading, natural ventilation, passive heating and cooling. Also, [19] identified four variables of bioclimatic architecture principles as sun shading devices, passive cooling system, thermal mass strategies and ventilation strategies.

What is a bioclimatic house? ›

A house that can go back to be part of the cycle of life.

What are the benefits of green building concept? ›

Green buildings can not only reduce or eliminate negative impacts on the environment, by using less water, energy or natural resources, but they can - in many cases - have a positive impact on the environment (at the building or city scales) by generating their own energy or increasing biodiversity.

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