Intelligent Buildings International

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Exploring design principles of biological and living

building envelopes: what can we learn from plant
cell walls?

Yangang Xing, Phil Jones, Maurice Bosch, Iain Donnison, Morwenna Spear &
Graham Ormondroyd

To cite this article: Yangang Xing, Phil Jones, Maurice Bosch, Iain Donnison, Morwenna Spear &
Graham Ormondroyd (2018) Exploring design principles of biological and living building envelopes:
what can we learn from plant cell walls?, Intelligent Buildings International, 10:2, 78-102, DOI:
10.1080/17508975.2017.1394808

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INTELLIGENT BUILDINGS INTERNATIONAL, 2018
VOL. 10, NO. 2, 78–102
https://doi.org/10.1080/17508975.2017.1394808

RESEARCH ARTICLE

Exploring design principles of biological and living building

envelopes: what can we learn from plant cell walls?
Yangang Xinga, Phil Jonesa, Maurice Boschb, Iain Donnisonb, Morwenna Spearc and
Graham Ormondroydc,d
a
Welsh School of Architecture, Cardiff University, Cardiff, UK; bInstitute of Biological, Environmental and Rural
Sciences (IBERS), Aberystwyth University, Aberystwyth, UK; cBioComposites Centre, Bangor University, Bangor, UK;
d
Department of Architecture and Civil Engineering, University of Bath, Bath, UK

ABSTRACT ARTICLE HISTORY

A number of innovations in building envelope technologies have been Received 9 August 2016
implemented recently, for example, to improve insulation and air Accepted 24 September 2017
tightness to reduce energy consumption. However, growing concern
KEYWORDS
over the embodied energy and carbon as well as resource depletion, is Adaptability; energy
beginning to impact on the design and implementation of existing and conservation measures;
novel building envelope technologies. Biomimicry is proposed as one ecological design; intelligent
approach to create buildings which are resilient to a changing climate, building; green building;
embedded in wider ecological systems, energy efficient and waste free. biomimicry; biomimetics;
However, the diversity of form and function in biological organisms and adaptable facade; plant cell
therefore potential applications for biomimicry, requires a holistic wall
approach spanning biology, materials science and architecture. It is
considered timely to re-examine opportunities to learn from nature,
including in the light of recent understanding of how plant form and
function are determined at the cellular levels. In this article, we call for a
systemic approach for the development of innovative biological and
living building envelopes. Plant cell walls are compared to building
envelopes. Key features of cell walls with the potential to inform the
development of design principles of biological and living building
envelopes are identified and discussed.

Introduction
The building envelope has been a significant element of human settlements since the rise of civiliza-
tion. It plays a dominant role in the exchange of heat and fresh air, provides views and daylight, and
protects the indoor environment and occupants against extremes of temperature, solar radiation,
water and wind. Vernacular building envelopes relied on local resources such as earth, timber, bam-
boo or stones. However modern building envelopes have utilized iron and steel over the last century,
and modified glass over the last few decades. Some materials have come into new eras, for example,
while the Romans used cement to make concrete, and to achieve radical new structures such as
domes, arches and vaults; modern Portland cement differs materially in several ways. For example,
the change to hydraulic lime in Portland cement in the eighteenth century increased industrial effi-
ciency of production (compared to Roman lime or gypsum alkali cements), a wider range of aggre-
gate is used depending on application, and the use of steel reinforced concrete to increase tensile and

CONTACT Yangang Xing [email protected], [email protected]

© 2017 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group
This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://
creativecommons.org/licenses/by-nc-nd/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the
original work is properly cited, and is not altered, transformed, or built upon in any way.
 INTELLIGENT BUILDINGS INTERNATIONAL 79

bending performance of the material has greatly extended the usefulness of concrete (Morgan 1977).
Timber structures have also entered a new era utilizing modern manufacture methods such as glue
lamination and cross-laminated timber (CLT) to allow new designs, long spans and tall timber build-
ings (Bjertnaes and Malo 2014; Epp 2016). In 1981, Davies proposed the concept of a ‘polyvalent
wall’ with multiple layers of glass materials which can generate enough energy for the building
(Davies 1981). Recently, building envelopes have been used to generate energy. Building integrated
photovoltaic (BIPV) approaches have been developed as more affordable building wall solutions
(Xing, Hewitt, and Griffiths 2011).
Recent changes in building regulations (such as Part L in the UK, and European directives such as
the Energy Performance of Buildings Directive (2010/31/EU), and the Energy Efficiency Directive
(2012/27/EU)) have promoted the use of more insulation materials and higher air-tightness of build-
ings (Jelle 2011; Xing, Hewitt, and Griffiths 2011). It is generally recognized that the operational
energy performance of both new and existing buildings will be improved dramatically through
the use of more insulation. However, what is often overlooked is the increased embodied energy
and carbon of many building materials, including synthetic insulation products – mineral wool
and plastic foams (Giesekam et al. 2014). Moreover, some researchers have argued that high insula-
tion may have adverse effects during summer in certain climate conditions (Stazi et al. 2015).
Resource intensive building design strategies (e.g. those containing over-sized insulation fabrics
and service engineering systems) have a significant deficiency when considering embodied energy
and carbon, which may lead to material depletion, unless a step change can be made in the sourcing
of building materials from renewable sources. The current resource depletion coupled with an
increasing demand for new buildings, due to rapid population growth and urbanization worldwide,
is leading to a number of environmental, social and economic issues. Current research efforts into
sustainable design practices are dominated by reductive approaches and hence their applicability
to a complete holistic design approach within architecture remains elusive (Gamage and Hyde 2012).
The climate is changing at an unprecedented rate, and may impose tremendous challenges for
future buildings (Xing, Lannon, and Eames 2013). Researchers have argued for a more holistic
approach to the design of buildings, considering all energy and sources of impacts (including
food and waste) (Vale and Vale 2010) using a whole ecosystem approach (Garcia-Holguera et al.
2016) as well as addressing societal changes (Xing 2013).The built environment has been considered
as a key element in ensuring the health and wellbeing of the population, reducing energy consump-
tion and carbon emissions. Therefore, new buildings need to be designed and constructed to be
adaptable and resilient to future climate change and fluctuations, with the existing building stock
retrofitted to achieve the same.
Through billions of years of evolution, nature has generated some remarkable systems and sub-
stances that have made life on earth what it is today. In order to remedy the destructive effects of
buildings, researchers have argued that it is important to create buildings resembling ecosystems
to increase resource efficiency and create cyclic resource loops (Benyus 2002; Pawlyn 2011; Gamage
and Hyde 2012; Zari, Pedersen Zari, and Zari 2015). Such learning from nature, or biomimicry, pro-
vides a platform to create a new generation of environmentally friendly and sustainable materials
and systems. It is therefore critical to change the views on the development of building technologies
and regulations based on nature’s wisdom to create buildings adaptable to the changing environment
and closely linked to ecosystems (Zari, Pedersen Zari, and Zari 2015).
Plants are constantly exposed to different environmental conditions. Being essentially sessile
organisms (i.e. fixed to the same habitat during their entire life cycle, with their only chance of dis-
persal through their seeds), plant survival is crucially dependent on adaptation to the changing
environmental conditions over a day and also between days, seasons and years. Recent advances
in plant science have uncovered the dynamic yet co-ordinated regulation of stress responses, pro-
cesses of growth, development and reproduction (Satake, Sakurai, and Kinoshita 2015). Buildings
can also be described as sessile (usually fixed to the same location). Both buildings and plants
have to be resilient and adaptable to the surrounding environments; therefore, there are potential
80 Y. XING ET AL.

opportunities to discover synergies between plants and buildings and identify potential biomimetic
solutions.
Ultimately, a plant’s adaptation to environmental stresses and conditions depends on responses
taking place at the cellular level. Plant cell walls are one of the defining differences between plant and
animal life forms, and the presence of these walls is a primary contributor to the evolution of land
plants as sessile organisms. The cell walls provide support, act as defensive layers, are conduits for
information, and are a source of signalling molecules and developmental cues. The cellular structure
of plants was discovered by Robert Hooke in 1665, and since this time the structure and function of
the cell walls have been studied in detail at cellular, genetic and molecular levels. Inspired by the
structure of plant cells, which is defined by their cell walls, a 3D-printed soft chair was created
using recyclable material (Martin 2014). In addition to the mechanical properties, plant biomass
also exhibits good thermal insulation properties. The cellular structure of cork has long been recog-
nized as a thermal and electrical insulator. The pith of many other plants can be used for similar
purposes, such as panels derived from hemp or flax shiv for lightweight structural or insulation
boards. Development of foamed insulation materials from either synthetic or bio-derived polymers
is an attempt to improve upon foams demonstrated in nature, by increasing thermal insulation
towards a conductivity of synthetic materials such as polyurethane foam or glass wool (25–
45 mWm−1K−1; Papadopoulos 2005), for example, tannin foams have now demonstrated thermal
conductivity of 75 mW m−1K−1 (Tondi et al. 2016). Other bio-based insulating materials rely on
natural fibres to provide loft for a low density batt or mattress of randomly aligned fibres (Kymalai-
nen and Sjoberg 2008). Not surprisingly, the use of cell wall biomass for developing energy efficient
and low cost construction materials is an emerging field in building construction and civil engineer-
ing (Vo and Navard 2016).
Plant cell walls have remarkable similarities with building envelopes in terms of providing struc-
tural support and protection from the external environment. However, there is a lack of research into
learning from plant cell walls to inform the philosophical debate of the development of resilient
building envelopes. The authors argue that building envelope design research and practices need
to learn from the adaptability and dynamic behaviour of plant cell walls. The key aim of this research
is to develop a holistic biomimetic approach to facilitate transformation of the building envelope
technologies. In this article, multiple functionalities of plant cell walls are reviewed; the analogy
between plant cell walls and building envelopes and existing efforts to develop bio-inspired building
envelopes technologies are identified; and a set of design principles for biomimicry transition is pre-
sented. The article concludes with opportunities and challenges for future development of living bio-
logical building envelopes.

A systemic biomimicry design framework: key components and a closed-loop

learning process
There is a rich and long history of gaining inspiration from nature for the design of practical
materials and systems. From the early nineteenth century, architectural designers and engineers
have started to imitate the forms, and develop new methods, analogous to the processes of growth
and evolution in nature and to apply aspects of biological thinking in innovative designs in general
(Steadman 2008). Researchers have also formed concepts around innovative biomimetic designs for
building applications (Vogel 2009; Vincent 2009). A number of terms have been used to describe the
process of learning from nature, and they are often used interchangeably, each with a slightly differ-
ent focus or starting point. For example, biomimicry promotes thinking of a building as a living
entity (Benyus 2002). Biomimetics, on other hand, a term coined by Otto Schmitt in the 1950s,
emphasizes the transfer of ideas and analogues from biology to technology (Schmitt 1969). A special
branch of biomimetics is phytomimetics which deals with plant-inspired materials, structures and
movements (Stahlberg 2009).
 INTELLIGENT BUILDINGS INTERNATIONAL 81

There are two general biomimetic design processes, that is, a top-down approach (technology
pull), and a bottom-up approach (biology push). The top-down approach starts from defining
human needs or a design problem and looking at ways in which ecosystems can provide solutions.
The bottom-up approach starts from identifying a particular behaviour or function of an ecosystem
and translating that into designs and products (Aziz and El sherif 2016). Researchers have argued
that the linear approaches (i.e. top-down or bottom-up) of biomimetics may only be sufficient if
the focus is on the abstraction of single functions (Knippers and Speck 2012). To design, construct
and maintain a building is a complex process which cuts across many disciplines and practices
requires systemic solutions.
Classifications of biomimetic design goals have also frequently focused on the outcomes obtained.
A commonly used classification is comprised of three main fields: structural biomimetics (i.e. con-
structions and materials in nature), procedural biomimetics (i.e. processes in nature) and informa-
tional biomimetics (i.e. principles of evolution and information transfer in nature) (French 2014;
Gebeshuber, Gruber, and Drack 2009). Mimicry of form or a single function are the most common
biomimetic principles reported, but these will have un-intended consequences and limited impact
for achieving the requirements of holistic building design. Mimicking biological processes and sys-
tems is harder to achieve, but will deliver greater impact (Garcia-Holguera et al. 2016).
We propose that the ideal biomimetic design process is to use an iterative closed-loop multi-dis-
ciplinary learning process (as presented in Figure 1). The key components of learning process
include: (1) to identify biological analogies as a foundation of future biomimicry design; (2) to estab-
lish novel design principles, and related technologies; (3) to develop and test prototypes. In order to
avoid following a linear and single function view, this iterative learning process (Figure 1) empha-
sizes the use of integrated biomimetic methods to stimulate biologists, architects and engineers to
develop fundamentally new research strategies and actions identifying new analogies and new design
principles and testing prototypes.

Inspirational biological analogies of living building envelopes – plant cell walls

One key element in the biomimicry research is to discover biological analogies of living building
envelopes so that to stimulate the creation of the prototypes. Multiple functions of plant cell walls
are explored in this article, which identify parallels for the future application of cell wall biomimetics
within architecture. A number of plant survival strategies to cope with changing environmental con-
ditions have been identified, such as adjustment of the timing of flowering in response to seasonal
changes in day length, to transportation dynamics of essential micronutrients (Satake, Sakurai, and
Kinoshita 2015). It is recognized that the multi-functionality of biological composite materials is
usually achieved based on a complex hierarchical architecture from nano- to macro-scale (Dunlop

Figure 1. A systemic biomimicry design framework.

82 Y. XING ET AL.

and Fratzl 2010). With the developments of micro-scale engineering in the physical sciences and
advances in micro biology, (Sarikaya et al. 2003), we propose that great potential lies in the learning
from plants at the micro levels (e.g. cellular level in this article), to inform future resilient building
design.
The analogy between plant cell walls and building envelopes might at first appear to rely only on
their common role as providers of protection and structural functions: strength, support, enclosing
spaces, and resistance to dynamic load. Indeed, plant tissues can provide structural integrity by
different routes. Examples include the structural optimization of cell wall components in xylem tis-
sue for load-bearing applications (Cave 1968), or the optimization of parenchyma tissue in shape and
cell wall structure to maximize control of turgor pressure, which provides a hydraulic function in
plant stem support (Wainwright 1970).
In addition to their structural role (strength of materials, control of turgor pressure and the
adhesion between cells which maintains plant integrity); plant cell walls also provide selective per-
meability of metabolites, enzymes and hormones, as well as facilitate cell to cell communication and
recognition, and response to stimuli. All cells have to maintain a certain rigidity to keep their shape
and to protect the elements inside. Although, plant cell walls and building envelopes have dramati-
cally different operational principles and mechanisms, they have remarkably similar key functions as
shown in Table 1. Here we argue that plant cell walls can provide an inspirational source of design
thinking to develop future bio-inspired building envelopes.

Structure, composite, form and functions

Plant cell walls: structure and composition
Our knowledge of plant cell walls is based on an in-depth understanding of its biosynthesis, structure
and molecular physiology. In his Micrographia, Robert Hooke discovered plant cells: more precisely,
Hooke had been viewing the cells in cork tissue and described them as an ‘infinite company of small
boxes’ saying that ‘these pores, or cells, were not very deep, but consisted of a great many little Boxes,
separated out of one continued long pore, by certain Diaphragms’ (Hooke 1665). Nehemiah Grew
and Marcello Malpighi carried out early studies on plant anatomy – revealing the diversity of
plant cell types, however understanding of the primary and secondary wall did not emerge until
the work of Kerr and Bailey in the twentieth century (Kerr and Bailey 1934). The plant cell wall

Table 1. Biological analogies: plant cell walls and building envelopes.

Analogy in key
functions Plant cell walls Building envelopes
Protection again . Provide a mechanical protection barrier against . Provide protection against external elements,
external biotic stresses (e.g. insects and pathogens) and helps such as wind, pollution, noise, solar radiation,
elements to protect against abiotic environmental stresses rain, and cold.
(e.g. wind, drought, heat, cold). . Maintain indoor climate.
. Separate interior of the cell from the exterior
environment.
. Prevent water loss.

Exchange of heat, . Enable transport of substances and information from . Conduits for plumbing, electrical and other
air and water the cell interior to the exterior and vice versa. services.
. Aid in diffusion of substances into and out of the cell. . Fenestration, ventilation ducting, and passive
. Design of pits for fluid flow and control of cavitation. air and moisture exchange.

To define shape . Give the cell a definite shape and structure. . Define space and function.
and space . Provide structural support. . Provide structural support and cultural
. Prevent the cell from rupturing due to turgor identify.
pressure.
 INTELLIGENT BUILDINGS INTERNATIONAL 83

is a highly complex structure that surrounds cells (as shown in Figure 2). It is located outside the cell
membrane and has a ‘skeletal’ role in supporting the shape and structure of the cell; a defining role in
differentiation of cell as one of the many cell types required to form the tissues and organs of a plant;
a protective role as an enclosure for each cell individually; and a transport role helping to form chan-
nels for the movement of fluid in the plant (Keegstra 2010). A segment of a stem cross-section in
maize shows the diversity of different cell types (Figure 3). Here sclerenchyma provide linear
strengthening to the relatively wide xylem and phloem cells in vascular tissue which are involved
in fluid transport. The parenchyma, with relatively shorter and broader cells provide a closed cell
foam maintaining the internal shape of the cylindrical stem to resist buckling (Alexander 2016).
Plant biomass consists predominantly of cell walls, typically 60–70% based on dry matter yield.
The cell wall consists of a sophisticated composite structure predominantly based on polysacchar-
ides, the most characteristic component being cellulose (the most abundant organic polymer on
earth). Microfibrils of crystalline cellulose, encapsulated in amorphous cellulose, are embedded in
a matrix of pectic and hemicellulosic polysaccharides (Keegstra 2010). Lignin, a heterogeneous aro-
matic and hydrophobic polymer that lacks a repeat structure (Boerjan, Ralph, and Baucher 2003),
may also be present in the cell wall of some plant tissues where it performs a bulking and an adhesive
role. Thus, the wall is assembled into an organized composite of microfibrils and matrix, linked
together by both covalent bonds and noncovalent bonds between macromolecules. It was recently
shown that xylan, the main hemicellulose polymer in secondary cell walls, slots together with cellu-
lose fibrils as a twofold helical screw (Simmons et al. 2016), revealing a previously unknown funda-
mental principle in the assembly of plant cell walls and improving our understanding of the
molecular cell wall architecture that makes very strong cell wall structures.
The cell wall composition, architecture, thickness and porosity varies from species to species, and
may also depend on cell type and developmental stage of the organism. Cell walls are a dynamic bio-
logical barrier that, together with the cell membrane (plasma membrane), separate the interior of all
cells from the outside environment. The plasma membrane, mostly composed of lipid molecules, is
selectively permeable to ions and organic molecules and controls the movement of substances into
and out of the cell (Furt, Simon-Plas, and Mongrand 2011).

Plant cell walls: composite structure and performance

The cell wall composition and structure give remarkable mechanical performance. Cell shape and
cell wall composition are optimized for the role of the cell within the plant. The arrangement of

Figure 2. Highly simplified model of the primary plant cell wall (based on McCann and Roberts 1991).
84 Y. XING ET AL.

Figure 3. Histochemically stained segment of a stem cross-section of maize.

Notes: Epi, epidermis; xy, xylem; par, parenchyma; phl, phloem; scl, sclerenchyma. Left hand side scale bar is 200 μm.

the four basic building blocks of plant cell walls: cellulose, hemicellulose, lignin and pectin, can result
in an exceptionally wide range of mechanical properties in plant tissues; and engineers have thus far
failed to achieve the same micro-structural control of composites as that exhibited by plant cell walls
(Gibson 2012).
In physical terms, the cell wall is a macro-molecular composite with some analogies to reinforced
concrete (Davison et al. 2013), with the chemical complexity and compact organization of cell walls
making it extremely resistant to deconstruction (Sarkar, Bosneaga, and Auer 2009). Thin crystalline
microfibrils of cellulose provide a reinforcing element within an amorphous cellulose and hemicel-
lulose matrix. Orientation of cellulose microfibrils within each layer of the cell wall is optimized. In
most plant cells, the primary role of the cell wall is to act as a pressure vessel (Wainwright 1970), with
the combined action of the cells acting as hydrostats to provide the elevation of the plant stem, leaves
or flower heads (Ennos 2012). In primary cell wall of parenchyma, where the role of the wall is to
provide resistance to hydrostatic pressure, the apparent amorphous alignment of microfibrils actu-
ally reflects optimal distribution to resist tension in all orientations, maintaining turgor pressure
within the cell. In tracheids and sclerenchema, where cells are elongated and secondary wall is sig-
nificantly thicker, the alignment of microfibrils helically around the axis of the cell provides optimal
resistance to longitudinal compressive forces, as well as enormous tensile strength. The multiple cell
wall layers, and their unique microfibril orientations (Figure 4) combine to provide the mechanical
properties of the composite cell wall structure.
Lignified tissues, such as the tracheids of the xylem in softwoods, have been well studied (Figure 4)
and the contribution of microfibril alignment within each layer of the wood cell wall to the mech-
anical properties of the woody tissue as a whole modelled (Mark 1967) to gain insight into the
 INTELLIGENT BUILDINGS INTERNATIONAL 85

Figure 4. Schematic of a tracheid in softwood xylem, indicating microfibril alignment in the primary and secondary cell wall layers.
Secondary cell wall comprises S1, S2 and S3 layers.

multiple functionalities of cell walls (Geirlinger et al. 2006; Cave 1968). The xylem provides a struc-
ture which is highly successful in resisting compressive loading, elevating the tree canopy tens of
metres into the air. The xylem within branches is adapted to resist bending loads, utilizing com-
pression wood (gymnosperms) or tension wood (angiosperms) in which the cell wall structure of
tissue below or above the pith (cells of the central portion) respectively has been altered to enhance
resistance to the named force. Despite this optimization, age, or extreme load can lead to failure,
however mechanisms such as formation of compression creases act to absorb energy, limiting the
extent of failure. Plants can also respond to their environment, especially when under stress, to
alter the amount of cell wall polymers (Gall et al. 2015) and also cell wall and whole stem structure,
for example, the production of tension wood or compression wood (Brereton et al. 2012). Such a
responsive structure could inspire the development of dynamic biological building envelopes.
Analogues for the cell wall design can be found in plywood and in synthetic fibre-reinforced com-
posites. The benefits of cross-laminating veneers of alternating grain direction were recognized by
the ancient Egyptians, and have been used in modern structures and aircraft, as well as plywood
itself. The design potential of angles other than 90° for the orientation of grain direction are well
explored in synthetic fibre composites, allowing curved panels, conical forms and complex cross-sec-
tions to be formed from continuous fibres. In the plant cell wall, the adaptability of microfibril angle
to contour around or reinforce apertures in cells provides numerous examples of bio-based design
optimization. Modern CLT has revisited the high strength and orthotropic character of plywood, in a
larger cross-section product suitable for construction of multi-storey buildings. The authors of the
article also suggest that a return to the fibre composite bioinspired design may lead to creation of
lightweight strong materials for use in walls, roofs and floors, potentially with combined secondary
functions such as ventilation, trunking or piping for underfloor heating.

Plant cell walls: form and function

The many plant tissues within the stem, the root, leaves or flowers, provide numerous examples of
differentiation in both form and function. Each tissue has a uniquely adapted assembly of cell wall
86 Y. XING ET AL.

components, utilizing rapid growth and self-assembly processes to differentiate the tissue for its role.
In each case, the alignment and optimization of location and angle of strong stiff cellulosic micro-
fibrils, and the composition of the matrix in which they are embedded (proportion of hemicellulose,
pectin, glycoprotein, and lignin) reflects the requirements of the cell wall in service.
Many structural tissues have cell walls which are optimized to resist turgor pressure. The firmness
of a fresh apple or carrot is very different to the lignified woody material of timber which relates to
lignification and thickness of cell wall secondary layers. Some other tissues have very specific roles in
which temporal changes in osmotic pressure achieve movement, such as the guard cells of stomata
on leaves, or the nastic movement in response to stimuli. The movement achieved is mainly gov-
erned by the cell shape and cell wall microfibril orientation. Thus while cell wall structure and
cell turbidity have a structural role, providing the upright stance of plants, they also govern move-
ment and will be considered further in the next section.

A brief overview of dynamic features of plant cell walls

Plant cell walls are highly dynamic and complex cellular structures supporting plant growth, devel-
opment, physiology and adaptation. Based on the brief overview of the plant cell wall properties, the
following three key features are introduced: porosity for filtration and communication, multi-func-
tional and dynamic materials, and biosynthesis process.

Porosity for smart filtration and communication

There are up to three major layers that can be distinguished in plant cell walls: the primary cell wall,
the secondary cell wall (where present), and the middle lamella. The middle lamella is the first layer
formed during cell division. This outermost layer is rich in pectin and joins together adjacent plant
cells. The thin, flexible and extensible primary cell wall is formed after the middle lamella while the
cell is growing and is the major textural component of plant-derived foods.
The primary cell wall is highly porous, and permits soluble factors to diffuse across the wall to
interact with receptors on the plant plasma membrane. Indeed, the primary wall contains up to
80% of its fresh weight as water. However, the cell wall is a selective filter that is more impermeable
than the matrices surrounding animal cells. With a pore size of 5–10 nm (Carpita et al. 1979), water
and ions can diffuse freely in cell walls, but diffusion of larger particles is reduced. The pectin net-
work appears to be a major player in dictating water content and porosity of the primary cell wall
(Mohnen, Bar-Peled, and Somerville 2008). Although secondary walls are typically much less
hydrated than primary walls, not much is known about their porosity, but lignin is thought to be
a key porosity gatekeeper in cells with lignified secondary cell walls. Many lignified tissues with sec-
ondary walls are designed for bulk flow, for example, xylem. In this case, the cell wall contains elab-
orate structures such as bordered pits, which regulate the flow of liquids and metabolites, while
providing some filtration or trapping effect against air bubbles or large impurities. The apertures
of the pits form during secondary cell wall development, sometimes with rearrangement of primary
cell wall components to increase the permeability in the desired direction, such as the margo strands
of bordered pits of conifer tracheids or the scalariform apertures in hardwood vessel cells (Wilson
and White 1986). This macro-scale flow is outside the scope of this section.
Even though plant cells are enclosed by a cell wall, cell to cell communication throughout plant
tissues is possible through structures called plasmodesmata, c. 50-nm-diameter plasma-membrane-
lined channels that connect adjacent cells through the cell-wall barrier (Ding, Itaya, and Woo
1999).The presence of plasmodesmata allows for a continuous cytoplasmic connection within
plant tissues called the symplast. There is a growing body of data showing associations of the cytos-
keleton, a complex network of actin filaments and microtubules, with plasmodesmata (Aaziz,
Dinant, and Epel 2001). Besides providing inner support for plant cells, the cytoskeleton, which
extends throughout the cytoplasm, is involved in intracellular trafficking and closely associated
with the plasma membrane.
 INTELLIGENT BUILDINGS INTERNATIONAL 87

Multiple functional and dynamic materials

In biology, the differences between material and system is blurred, and biological materials are often
part of a structural system. In the past, the plant cell wall was often viewed as an inert and static
exoskeleton. It is now recognized as a highly dynamic structure that, besides providing mechanical
support, needs to respond to various environmental and developmental cues and fulfils important
functions in signalling events, defence against biotic and abiotic stresses, and growth (Keegstra
2010). In addition, the structural shape of the plant is not solely reliant on the shape of its constituent
cells, but able to grow, flex, open and close flowers or leaves, and to adjust angle or orientation to
maximize sunlight. These functions are achieved by response to hormones, or following circadian
rhythms, or the use of osmosis to alter turgor pressure in selected tissues. The dynamic nature of
the wall, needing to be responsive and adaptable to normal processes of growth as well as to stresses
such as wounding, attack from pathogens and mechanical stimuli, requires sensing, signalling and
feedback mechanisms. The emerging view on the plant cell wall is one of a dynamic and responsive
structure that exists as part of a continuum with the plasma membrane and cytoskeleton (Hum-
phrey, Bonetta, and Goring. 2007; Baluška et al. 2003), although the exact linkages between these
three components are still not well defined (Liu, Persson, and Zhang 2015).
In non-lignified plant tissues, it is the internal pressure of the cell contents that allows plants to
maintain their upright stance. The cell wall enables plant cells to develop high turgor pressure (typi-
cally 0.3–1 MPa), important for the structural stability of the cells within plant tissues (Cosgrove
2009). The turgor pressure also influences the water relations and water economy of plants; the
loss of turgor pressure, that is, when the rate of loss of water from the plant is greater than the
absorption of water in the plant, for instance, due to drought stress, causes wilting. To resist internal
hydrostatic pressure, the microfibril alignment in the primary cell wall is optimized to achieve hoop
strength of the cell (Wainwright 1970). This is different to the load-bearing role of the secondary cell
wall discussed in the section ‘Structure, composite, form and functions’. This combined action of the
cells under hydrostatic pressure within a closed cell foam can provide significant hydraulic support
to plant tissues. In addition, control of hydrostatic pressure by osmosis allows response to stimuli
and nastic movements as mentioned above, with leaf angle or flower head tilting being a result of
short-term alterations in the turgor pressure. The touch response of Mimosa pudica is a well-
known example (Volkov et al. 2010). Here the shape and location of the parenchyma cells within
pulvini govern the range of movement, and the electrical signalling mechanism allows rapid response
by the leaflets. Tropic movement, by adjustment of cell growth in response to light or gravity, also
overcomes some of the limitations of the sessile nature of plants, allowing growth into adjacent
spaces as a response to changes in the canopy or competitor plants. These responsive structures pro-
vide inspiration for mechanical devices and actuators within buildings, as will be discussed in the
section ‘Key novel design principles and related attempts’.

Biosynthesis process and programmed cell death

Self-assembly allows plants to accommodate the changing needs of the growing plant cells and the
broad variety of cell shapes and functions. Being dynamic structures which are continuously syn-
thesized and remodelled during plant development, it is probably not surprising that the biosynthesis
of plant cell walls is a complex and highly regulated process (Guerriero, Hausman, and Cai 2014). To
illustrate this, plants invest a large proportion of their genes (∼10%) in the biosynthesis and remo-
delling of the cell wall (McCann and Carpita 2015).
Plant cell walls also contain structural proteins, enzymes, and other materials that can modify the
physical and chemical properties of the cell wall. It has been estimated that more than 65 different
enzymes are required to synthesize the pectic polysaccharides known to exist in plant cells (Harholt,
Suttangkakul, and Scheller 2010). Figure 5 shows an example of a highly specialized plant cell, the
pollen tube and highlights the polarized pollen tube growth process and shows the thickening of
the cell wall at the apex induced by exposing the pollen tube to a particular enzyme that changes
the mechanical properties (Bosch, Cheung, and Hepler 2005).
88 Y. XING ET AL.

Figure 5. The pollen tube, a highly specialized plant cell with a dynamic cell wall at the apex. Microscopy images showing a time
series of a Lilium formosanum pollen tube growing in in-vitro growth medium.
Notes: Numbers represent minutes after addition of an enzyme (pectin methylesterase) to the growth medium that changes the cell wall properties,
leading to the arrest of pollen tube tip-growth. Scale bar = 10 μm. Adapted from Bosch, Cheung and Hepler (2005) and used with permission.

For plants to develop properly and survive, including in response to environmental challenges,
they need to be able to make radical changes including to re-design and re-engineer their basic struc-
ture. Programmed cell death (PCD) provides an important response strategy to various internal and
external cues (Lam 2004). PCD is a highly regulated process for the selective dismantling of
unwanted cells and is essential for plant growth and survival as it plays a key role in embryo devel-
opment, formation and maturation of many cell types and tissues, and plant reaction/adaptation to
environmental conditions. For example, PCD as a final stage of differentiation in xylem tracheary
elements results in a continuous system of adjoining hollow cells that function in water/solute trans-
port. Here PCD accompanies the lignification of the cell wall, leaving dead tracheid cells as structural
tissue optimized for fluid flow. The suicide of a cell through PCD involves the execution of a geneti-
cally encoded and actively controlled sequence of steps.
Our current understanding of PCD in plants is largely shaped by research on animal PCD, par-
ticularly apoptosis. However, it is often forgotten that the concept of PCD originated from plants
(van Doorn et al. 2011). Although plant and animal PCDs share numerous characteristics (for
instance, nuclear DNA degradation), several differences exist. The presence of a thick cell wall dic-
tates that plant cells are not phagocytic (engulfment of cell corpses by another cell) and that corpse
clearance is a cell autonomous process in plants. The dying cell synthesizes substances, including
lytic enzymes, to break itself down and places them in the vacuole that ruptures as the cell dies
(van Doorn et al. 2011). Within buildings the potential to form structural material or conduits for
services in situ during construction would mimic the plant PCD mechanisms, whereas design for
deconstruction at end of life requires a more radical animal PCD approach.
In summary, the brief overview of the plant cell wall characteristics demonstrates a number of
interesting properties that merits further exploration for the design of bio-inspired building envel-
opes, further discussed in the following section.

Key novel design principles and related attempts

Based on the above review of plant cell wall characteristics, the following three key novel design prin-
ciples of the biological and living building envelope are proposed: permeable, shape changing, and
biosynthesis process. Nevertheless, learning from plant cell walls to inform the development of future
biological building envelopes is in its infancy. However, a number of related attempts have been
made to develop bio-inspired building envelopes (as is summarized in Table 2). Those existing
attempts are not directly linked to the learning from the plant cell walls, but it may promote discus-
sion and shed light on the future development of a potential technical pathway (as illustrated in
Figure 6) to incorporate learning from plant cell walls into the design of biological building
envelopes.
 INTELLIGENT BUILDINGS INTERNATIONAL 89

Table 2. Related attempts ad demonstration examples of living building design principles.

Design principles Building examples Technologies References
Permeable and Cellular envelopes Cellular structures serve as both a Grobman (2013)
multiple Dynamic insulation materials structure and a barrier Taylor and Imbabi (1999)
functional Transpired solar collectors Porous façade to allow heat exchange Love et al. (2014); Shukla
Porous reflective cool roof Perforated steel skins to provide heat et al. (2012)
‘Polyvalent’ wall exchange and air flows Craig et al. (2008)
Air filtration to improve air Porous roof structures with orientated Davies (1991)
quality holes which can re-radiate heat Stefanowski et al. (2015)
Multiple layers of glass materials and PV
can generate energy
Porous materials with capacity to absorb
vapours and VOCs
Adaptable shape Hygromorphic materials Response driven by the shrinkage and Holstov, Bridgens, and
changing Mechanical responsive swelling of wood triggered by moisture Farmer (2015)
facades changes. Loonen et al. (2013);
Kinetic skin Dynamic shape changing facades with Christoforou et al. (2013);
Shape changing materials mechanical actuators Kirkegaard (2011)
Dynamic light transmittance High tensile strength with low bending Knippers and Speck (2012)
glazing stiffness of lamellas allowing elastic Fiorito et al. (2016); Bar-
deformations Cohen (2005)
Shape changing polymers, alloys or Mardaljevic, Waskett, and
hybrid materials, for example, EAPs, Painter (2015); Aldawoud
PZTs (2013)
Electrochromic glazing materials
Biosynthesis process Vegetated buildings Green roofs, green walls and ‘tree houses’ Xing, Jones, and Donnison
Solidified granular materials Cementation of sand dunes to create a (2017)
Grow buildings using network of sand dunes to prevent Larsson (2011)
multiple functional desertification Imhof and Gruber (2015,
biomaterials Mycelium building materials blocks and 2017); Benjamin (2016)
Cellophane house slime mould to locate optimal space or Kieran and Timberlake (2008)
routes
Using discrete components in reversible
process

Permeable and multiple functional building envelopes

Key permeable and multiple functional features of plant cell walls
The permeability of plant cell walls plays vital roles for filtration, sensing and communication,
whereas, permeability of built walls is rarely considered in building design. The modern use of moist-
ure barriers and other membranes has reduced permeability of structures overall. On the other hand,
the plant cell wall together with the plasma membrane, the latter mostly composed of lipid

Figure 6. Transformation pathway.

90 Y. XING ET AL.

molecules, is selectively permeable to ions and organic molecules and controls the movement of sub-
stances into and out of the cell (Furt, Simon-Plas, and Mongrand 2011). Plasmodesmata in the plant
cell wall allow cell to cell communication, connecting with the cytoplasm to maintain a continuous
symplastic pathway, as discussed earlier in the section ‘Porosity for smart filtration and
communication’.

Related attempts in creating permeable building envelopes

Vincent (2009) argued that ‘functional’ form is one of the most important parameters in biomimetic
design. There have been few attempts to develop porous building envelopes. Inspired by cellular/
spongy envelopes in nature, researchers have been developing a framework for creating new
forms of building envelopes based on a complex cellular or sponge-like geometry and preliminary
design experiments in which cellular envelopes serve as both a structure and a barrier (Grobman
2013). Taylor and Imbabi (1999) explored ideas of a porous façade to allow heat exchange as
dynamic insulation. Craig et al. (2008) proposed porous roof structures with orientated holes
which can re-radiate heat to the night sky and control temperature of a building. The design of
the apertures was a critical factor, with not only orientation but also the pore dimensions being
selected to allow loss of long wavelength infrared radiation while resisting convective heat loss. Sev-
eral researchers argued that transpired solar collectors (perforated steel skins) can provide a number
of functionalities, such as heating spaces, providing warm ventilation air, and supplying domestic hot
water in summer (Love et al. 2014; Shukla et al. 2012).
Soar (2015) argued that traditional and natural materials (such as earth, wood and straw) have
complex porous structures and are ‘intelligent’ in responding to the natural environment. For
example, clay, whether used in adobe construction or termite mounds, responds to water vapour
in remarkable ways and this interaction makes them natural phase change materials. Rapid develop-
ments in wood modification over the past decade has led to the improvement of dimensional stab-
ility, decay resistance or strength of timber, which is now also being developed in wood based
composite panels (Ormondroyd, Spear, and Curling 2015). However, new materials and designs
for the skin and core of structural and insulated panels need to be developed to improve their effi-
ciency and robustness (Panjehpour et al. 2013; Chen and Hao 2015).
Sorption of vapours and volatile organic compounds (VOCs) from the environment is an impor-
tant role in the mechanical ventilation or heat recovery systems of buildings, maintaining indoor air
quality. This is frequently achieved with filters that show high sorption for pollutants and odour mol-
ecules. The ability of certain natural building materials to scavenge VOCs from the atmosphere has
been demonstrated (Mansour et al. 2016; Stefanowski, Curling, and Ormondroyd 2016). Work has
been undertaken on the assessment of nut shells for their ability to be used in wall panels to improve
indoor air quality by the sequestering of VOCs from the atmosphere (Stefanowski et al. 2015).
Further work to incorporate scavengers within the surfaces of building products, or deliver these
within paints and coatings, will provide a passive control of odour and removal of undesirable VOCs.

Adaptable shape changing

In biology, the differences between material and system is blurred, and biological materials are often
part of a structural system. In the past, the plant cell wall was often viewed as an inert and static
exoskeleton. It is now recognized as a highly dynamic structure that, besides providing mechanical
support, needs to respond to various environmental and developmental cues and fulfils important
functions in signalling events, defence against biotic and abiotic stresses, and growth (Keegstra 2010).

Key adaptable shape changing features of plant cell wall

The plant form (or plant shape) is the result of the combination of tissue types, which in turn are
defined by adaptations in developing cell wall architecture during cell differentiation. Some of
these features allow cells to be responsive during the life of the plant, for example, the opening
 INTELLIGENT BUILDINGS INTERNATIONAL 91

and closing of stomata is regulated by changes in turgor pressure. Others such as nastic and tropic
responses may also result in alteration in cell wall to result in growth or movement towards or away
from the stimulus.
A number of plant survival strategies to cope with changing environmental conditions have been
identified, such as adjustment of the timing of flowering in response to seasonal changes in day
length, to transportation dynamics of essential micronutrients (Satake, Sakurai, and Kinoshita
2015). Plant cell walls are highly dynamic structures offering dynamic and multiple functionality.
Existing building envelopes are static and cannot adequately respond or connect to the surrounding
ecological systems. There are two types of plant movements: one group is water-driven movements
(growth, swelling/shrinking of cell wall) and the other group uses elastic instabilities to amplify the
capacity to move. The second group includes the use of shape as well as material structure to create,
for example, the snap closure of a Venus fly trap, or the explosive fracture of seed pods which pro-
vides a catapult action aiding dispersal (Skotheim and Mahadevan 2005). Researchers have argued
that nastic structures of plants and their reversible movements represent a recurrent model to be
mimicked (Guo et al. 2015; Fiorito et al. 2016). Plants do not directly rely on metabolism to produce
motion and are able to produce ‘muscle-less’ movement and stiffness which offers a means of achiev-
ing a significantly advanced architectural material system (Jeronimidis 2009). These systems are
particularly suitable as passive actuation which does not require active metabolism (Forterre 2013;
Guo et al. 2015).
In each case, the alignment and optimization of location and angle of strong stiff cellulosic micro-
fibrils, and the composition of the matrix in which they are embedded (proportion of hemicellulose,
pectin, glycoprotein, and lignin) reflects the requirements of the cell wall in service. Significant ten-
sile or compressive forces can be achieved by control of the angle of winding within cylindrical cells,
as described by Fratzl, Elbaum, and Burgert (2008).

Related attempts to create adaptable shape changing building envelopes

There have been a number of attempts to create adaptable building envelopes. Stazi et al. (2015)
examined external insulation layers that can be sealed in wintertime and ventilated in summer to
improve energy efficiency of the buildings. Recently, a number of researchers have advocated mova-
ble shading devices powered by electrical actuators to reduce energy consumptions in buildings
(Loonen et al. 2013; Christoforou et al. 2013; Kirkegaard 2011). However, most of the shape chan-
ging shells rely on mechanical actuators. Designs using shape memory alloy actuators for façade con-
trol have been discussed (Pesenti, Masea, and Fiorito 2015). Developments of hygromorphic or
temperature responsive actuators could further improve passive building climate regulation. As
an alternative strategy, researchers have demonstrated that electrochromic glazing can moderate
solar heat gains and reduce, or even eliminate, the need for moveable internal shading (e.g. venetian
blinds) and fixed external blinds (e.g. brise-soleil) (Aldawoud 2013; Mardaljevic, Waskett, and Pain-
ter 2015).
One potential application for responsive sensors within buildings would be in natural ventilation
systems. A theoretical precedent can be taken from the timber wall structures of boat houses in
Norway, in which the natural movement of plain sawn timber boards is harnessed to provide a natu-
rally opening louvre driven by the shrinkage and swelling of wood triggered by moisture changes.
Several research groups have recently developed hygromophic (moisture-responsive) materials uti-
lizing the anisotropic response of the wood cell wall to moisture or humidity uptake (Holstov,
Bridgens, and Farmer 2015). This utilizes the difference in swelling between different orientations
of wood veneers to create a flat or a curved section, using a two-layer structure in which the
grain orientation of the wood is differently aligned. Here the significantly greater swelling of
wood in its transverse direction than its longitudinal direction leads to differential swelling, and
induces curvature in the component. The correct selection of wood growth ring orientation allows
the board to flex to an open state in summer, in periods of low humidity, and to close due to moisture
uptake within the wood in winter, relating to high humidity. The process is governed by the lateral
92 Y. XING ET AL.

swelling of the wood being greater in the direction tangential to the growth rings than the radial
direction, producing a cupping effect in the plank. Careful selection of plank orientation during
installation allows the distortion to provide ventilation at the preferred time of year, and has been
likened to the mechanism of a pine cone opening to release seeds when dry.
The difference between tangential and radial orientations can also be harnessed for more subtle
effects. Complex forms can be created, inducing torsion rather than curvature in a mode which better
models the seed pod movement mechanism (Ionov 2013). Examples of hygromorphic materials
directly inspired by the pine cone, have been used in adaptive facades of prototype buildings,
where they introduce passively controlled permeability (Menges and Reichert 2012), and have poten-
tial for use in other areas of engineering, design and medicine. The same principle has been used with
hydrogels, polyelectrolyte layers and conducting polymers to create hygromorphic or thermally
responsive actuators (Ionov 2013). The microstructure and orientation of fibrils within layers of
the hygromorphic material governs the direction and magnitude of the response.
Researchers have attempted to develop new types of composite containing an integrated high
density of small sensors that would enable sensing without compromising the structural integrity
(Sagi 2005). Prototypes using materials with both sensors and actuators (e.g. alloys, polymers or
hybrids) that respond to an external stimulus to provide shading effects have been reviewed (Fiorito
et al. 2016). However, there are a number of challenges associated with sensorized composites,
including electronics, mechanical integration, and data management. Kinetic skins are developed
utilizing high tensile strength combined to a low bending stiffness of 108 lamellas allowing large elas-
tic deformations (Knippers and Speck 2012). The variable lateral openings of the kinetic skins are
used to control the lightning conditions of the interior spaces.
Conceptual models have been discussed to develop bio-sensing systems (Biggins, Hiltz, and Kus-
terbeck 2011). While research into developing bio-inspired sensing systems is in its infancy, several
examples can be found where biomimetics has contributed to actuator development. New hybrid
materials (bio and non-bio materials) are being developed. For example, a number of shape changing
materials have been reviewed, such as electroactive polymers (EAPs), piezo-electrical material PZTs
and shape memory alloys, polymers or hybrid materials, which have been used either as actuators or
sensors (Fiorito et al. 2016). However, the materials are still limited in their ability to generate suffi-
cient force to perform significant tasks, such as lifting heavy objects (Bar-Cohen 2005). Other new
materials have been developed based on nanotechnologies to offer emerging functionalities, such as a
prototype of new biosynthetic materials that function as self-healing membranes (Speck et al. 2006)
and self-cleaning photocatalytic building materials (Pinho, Rojas, and Mosquera 2014).

The biosynthesis process of living building envelopes

Key biosynthesis features of plant cell wall: growth and disassembly
The plant cell wall forms an excellent example of how nature can use a few widespread natural con-
stituents (usually C, H, O and N) to tailor molecules of diverse structures performing a wide variety
of functions. Plant cells are continuously synthesized and remodelled during plant development to
accommodate growth and cell differentiation. In addition, PCD allows plants to respond the chan-
ging requirements by terminating the function of cells once their role has been accomplished. Both
concepts (growth and programmed death) can be transferred to the built environment.

Related examples of growing biological building envelopes

One good example of biological building envelopes is the green roofs/walls, which can be installed on
most of existing buildings (Xing, Jones, and Donnison 2017). Researchers have also used traditional
‘pleaching or grafting’ techniques (Seymour 1976), which involve interweaving branches (living and
dead) through a hedge or steel structure to create prototype green façades, for example, ‘tree houses’
(Joachim 2016), or ‘Baubotanik’ which combine steel scaffolding with living plants (Ludwig 2016).
However, there is a need to improve the design and maintenance (e.g. choices of plants, substrates
 INTELLIGENT BUILDINGS INTERNATIONAL 93

and configuration) of green building envelopes to maximize the potential benefits (such as thermal
comfort, biodiversity). Researchers have also argued for a radical shift in construction, towards the
localized cementation of granular materials, for example, creation of a network of solidified sand
dunes to prevent desertification (Larsson 2011). The growing of biological building envelopes has
great potential in the future to further reduce or de-couple from consumption of fossil fuels based
resources.
Compared to steel and cement, biological materials are often lightweight and can be generated at
ambient temperature. Experiments have been set up to utilize mycelium (a fast-growing vegetative
part of a fungus) as a scaffolding structure to consolidate fragmented matter producing solid building
materials out of waste products from wood (Imhof and Gruber 2015; Benjamin 2016; The 3 Foragers
2013). Gruber and Imhof (2017) also introduced an experiment using slime moulds (a single cell
organism) to show its space path-finding capacity. Researchers have proposed that architectural
‘organ’ systems might act as hubs of bio/chemical activity, flow and transformation (Spiller and
Armstrong 2011; Armstrong 2016). Nevertheless, research activities exploring the concepts of grow-
ing buildings as a biological organisms are in very early stages (Gruber and Imhof 2017).

Related attempts in programmed demolition and retrofitting of building envelopes

Within buildings the potential to form structural material or conduits for services in situ during con-
struction would mimic the plant PCD mechanisms, whereas design for deconstruction at end of life
requires a more radical animal PCD approach. PCD in the biological kingdom can inform design for
deconstruction, or for development of adaptable building spaces. The phagocytosis process may
inspire design of building components which are readily removed or re-located, or components
which are easily recycled, industrially composted or suitable for recovery of monomer for new
materials production. Waste materials generated from building construction and demolition have
become a great challenge to sustainable urban development (Xing et al. 2009; Xing, Lannon, and
Eames 2014). Learning from PCD which serves fundamental functions during an organism’s life-
cycle, new perspectives in constructing regenerative building envelopes and developing sustainable
demolition strategies can be developed. New research activities are needed to develop programmed
building demolition or retrofitting as a part of a biosynthesis process. The cellophane house concept
demonstrated by Kieran Timberlake at the Museum of Modern Art centred around this shift away
from permanence in buildings – with multiple discrete components within panels held by quickly
reversible processes. The integrity, reusability and upgradability of the components was central to
the design (Kieran and Timberlake 2008).

A basic conceptual prototype, challenges and opportunities

In the light of the parallels between plant cell walls and the emerging architectural concepts and
available bio-inspired materials, the challenge facing architects and engineers is to combine these
technologies and concepts into a holistic solution. The concepts of wall permeability and regulation
of interior conditions by passive motion offer potential for new structures. The authors now consider
a conceptual design to integrate these features within a functional unit.
A basic conceptual design of a biological dome constructed using biological composite panels is
presented in Figure 7 to illustrate that the three key design principles (i.e. permeability, shape chan-
ging, and biosynthesis process) can be realized through the changes of physical (e.g. hygrothermal or
electrochemical) conditions of the panels which trigger the changes of the opening size, heat transfer
coefficients, lighting transmittance, air exchanges, and solar gains to optimize energy performance of
the buildings. The biological panels of this prototype can be generated using biomass waste to reduce
the embodied energy. As shown in Figure 7, each of bio-panels can be changed individually.
However, there is also a trade-off in the potential shape changing behaviours. For example, higher
daylight penetration can reduce electric lighting energy consumption but may also increase building
heating or cooling energy consumptions. Furthermore, different bio-panels may require different
94 Y. XING ET AL.

Figure 7. An illustration of the bio-dome concept.

behaviours during different time of the day in different climate zones. In order to determine the opti-
mal shape changing behaviour of each panes in the bio-dome, a preliminary theoretical optimization
algorithm was developed as part of an on-going project using computer simulations to develop most
efficient adaptable bio-dome buildings designs. Based on the theoretical optimization algorithm, key
theoretical physical characteristics of ideal materials can also be identified. The objective function of
this model is to minimize total energy (operational and embodied) consumption.
The optimization objective is to identify:


Min energy consumption = {heating, cooling, ventilation and lighting}


+ {embodied energy}.

Optimization constraints are the maximum and minimum theoretical physical limitation of each bio
panels, such as heat transfer coefficient, air infiltration rate, lighting transmittance, and embodied
energy of the bio-panel:
Umin ≤ Ui (Heat transfer coefficients U - value of bio - panel i) ≤ Umax ,

ACHmin ≤ ACHi (Air infiltration value of bio - panel i) ≤ ACHmax ,

LTmin ≤ LTi (Light transmittance value of bio - panel i) ≤ LTmax ,

EEmin ≤ EEi (Embodied energy of the bio - panel i) ≤ EEmax .

A number of high performance passive engineering materials (as listed in Table 2) can fulfil some of
the design goals of the living walls. For example, porous and reflective materials can provide dynamic
insulation (Taylor and Imbabi 1999; Love et al. 2014, Craig et al. 2008) and improve air quality (Ste-
fanowski et al. 2015); hygromorphic materials (Holstov, Bridgens, and Farmer 2015), elastic panels
(Knippers and Speck 2012), mechanic actuators (Loonen et al. 2013) and electrochromic materials
(Mardaljevic, Waskett, and Painter 2015) can regulate air flows and control daylight penetration and
solar gain. The development and integration of shape morphing panels with adjustable shading via
hygrothermal, EAPs, photochromic glazing and mycelium materials provide promising alternative
materials, however some of these features conflict with one another – requiring innovative design
to combine the elements effectively. To be able to optimize the system for solar gain, ventilation,
thermal comfort and day lighting simultaneously a hybrid system must draw on more than one tech-
nology. This will require continued research to optimize effects on interior climate and development
 INTELLIGENT BUILDINGS INTERNATIONAL 95

of working prototypes. By identifying a conceptual structure and model system, it is possible to

address the mutual interaction of competing insulation, permeability, daylighting and shading
requirements. Furthermore, the future integration of self-assembly concepts and design for decon-
struction requires a step change in building design theories and philosophies.
Building designs have generally evolved to have solid, impermeable envelopes and permeability is
currently facilitated though openings and ducts for perfectly valid reasons; perhaps controlling per-
meability though mechanical means is easier and more controllable than a porous façade. However,
as discussed previously current construction practices rely on mineral and fossil fuels based materials
which are finite resources. Renewable biological materials which can be generated based on circular
bio-economy may eventually replace current materials and building forms. Therefore, in the post-
carbon societies (Xing 2013), we will have to design, use and maintain the buildings differently.
Future research investigating multiple functionalities of the materials at different scales in the
hierarchy (from nano to macro and tissue and whole plants) is needed to develop and scale-up bio-
mimetic design principles for district or urban planning. However, allometric scaling laws need to be
considered (West, Brown, and Enquist 1997). Biological role models very often have to be scaled up
to a much larger size than their original size, which leads to difficulty for functional requirements.
Biological building envelopes might need to be constructed somewhat differently from the shape or
form of the original plant cell walls. The biomimetic structure or material must therefore address
differences relating to the change of target property which may be governed by a power law or allo-
metric scaling rather than relationship to the altered dimension.
Integration of advanced wall constructions based on plant cell wall inspired materials and build-
ing control concepts requires further study, and building physics models need to be created in the
context of architectural geometries for detailed analysis of energy flux and in service performance.
The advances in architectural software tools will allow researchers to analyse materials and designs,
and provide a visualization method to illustrate dynamic biological systems. Ultimately advanced
computer tools, micro-robots and micro-mechanical systems, laser cutting, 3D printing and other
digital design technologies can help researchers and practitioners to create and investigate future bio-
logical and living structures.
It is not a trivial task to establish different requirements, objectives and goals in the natural world
for biomimicry research and practice. Clearly implementing biomimetic design methods can be dif-
ficult and involves a long period of adaptation, depending on many factors such as technology matu-
ration, social acceptance and economic efficiency. The application of biomimetics in industrial
design and product development requires a process of adaptation to traditional methods through
simple models and a learning system. The key related technological solutions presented in this article
are far from exhaustive. Future research will be needed to establish the best practices and assessment
standards for implementation. Moreover, implementation of the new design principles needs to be
supported by policymaking and community engagement to gain maximum benefit from sustainable
and bio-inspired designs in buildings.

Conclusion
In this article, the authors have identified a series of analogies between the plant cell wall and the build-
ing envelope. The comparison of the static structural functions and the dynamic role of the wall during
the lifespan of the cell reflects the vital functions, including definition of space, osmotic and physical
protection, selective permeability barrier, immobilized enzyme support and cell–cell communication,
recognition and adhesion, and PCD. Bringing together the disciplines of architectural design, plant
biology and materials science, in this article, we promote the concept of biological building envelopes
based on studies of the fundamental structure of plant cell wall. It is pertinent to identify opportunities
to enable people from different disciplines to work together, to identify challenges and possible res-
olutions. This article explores what building designers can learn from plant cell walls at a cellular
level and from evolutionary concepts for the transformative design of building envelopes.
96 Y. XING ET AL.

Holistic biomimetics research is more than just a one-way knowledge transfer from biology to
technology. There is also a valuable contribution to be made by engineers and designers to help biol-
ogists to resolve the design complexity and identify operational principles within and behind the
natural world. In this holistic manner, the interdisciplinary research can bring mutual benefits to
ecosystems, as well as to the development of diverse research disciplines and practices, such as
biology, architecture, materials sciences and engineering. The cell wall composition, architecture,
thickness and porosity varies from species to species, and may also depend on cell type and devel-
opmental stage of the organism. Plant cell walls are highly complex structures (Rafelski and Marshall
2008) and there is a lack of research activities investigating energy and mass transfer between cells
and their environment. Thermal and mass transfer is a key research area established by building phy-
sics professions. Therefore, building physics tools may be able to contribute to the future develop-
ment of plant cell wall studies in order to inform future biomimicry designs.
Several areas of bio-inspired design – either materials harnessing mechanisms demonstrated in
plants, or the use of materials to achieve passive regulation of interior climate have been highlighted.
Rapid research progress is underway within architecture, as typified by the adaptive building facades,
use of bio-based materials, energy harvesting and selective energy re-release or optimization of pas-
sive ventilation. This article can present only a selection of highlights in this sphere in order to draw
attention to future challenges. Key principles include the use of self-assembly in creation of cell walls
with optimized fibril alignment to form composites with multiple functionality. The optimized pore
dimensions allowing communication and filtration, and the use of PCD to create rigid structures for
fluid transport. In the plant limited resources are used with maximum efficiency. The principles of
nastic movements in plants are particularly discussed with relevance to passive control of interior
climate and occupant comfort in buildings. The authors hope that building researchers can appreci-
ate the complexity of plant cell walls and promote activities to seek the key features of future biologi-
cal building envelopes and to develop the necessary technical pathways.
Current building practices are having an adverse effect on nature, for example, depleting
resources, reducing biodiversity, and generating pollution and waste. The authors argue that there
is a need to re-examine the fundamental concept of the building envelope which currently only
serves as a barrier, and is not connected with its surrounding ecosystems and there is a need to
develop new biologically inspired intelligent systems for buildings to support the processes of life
rather than relying on fossil fuel-based construction process. Furthermore, fundamental changes
of the design philosophies and technologies are needed to develop the next generation of building
envelopes. In order to transform existing static building envelopes to biological, intelligent and living
building envelopes, building designers need to take the lead in proposing new frameworks, leading to
more ambitious architectural practices to develop ecologically responsive buildings as guardians for
their inhabitants.

Disclosure statement
No potential conflict of interest was reported by the authors.

Funding
The Authors acknowledge the financial support of the Welsh Government and Higher Education Funding Council for
WalesFunding Council for Wales through the Sêr Cymru National Research Network for Low Carbon, Energy and
Environment.

Notes on contributors
Dr Yangang Xing is a building physicist investigating the interactions between plants and architecture (e.g. the nexus
of green infrastructure and urban heat islands/air quality, energy crops, organic building materials, biomimicry and
food) through development and innovative applications of building physics research tools and, in a broader context,
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systemic modelling and assessment of future post-carbon building environment. Yangang is devoted to identify and
develop new research applications to uncap the full potentials of multidisciplinary collaboration to tackle the grand
challenges facing society on the horizon. Yangang completed his PhD in Dynamic whole system simulation for sus-
tainability planning, and has done EPSRC-funded research focusing on methodological and technical innovations sup-
porting building and urban sustainability.
Prof. Phil Jones is a building physicist with an interest in low-energy, low-carbon, and sustainable design in the built
environment. His research interests include the development of computer models for energy and environmental pre-
diction, urban scale sustainability, research through design, and building energy and environmental monitoring. He
has worked on the use of plants to improve the micro-climate around buildings, and simulating their effect on reducing
external temperatures. He is Chair of the Board of Directors of Warm Wales, a community interest company formed to
install energy efficiency measures to existing fuel poor housing in Wales. He chairs the Welsh Government’s Building
Regulation Advisory Committee.
Dr Maurice Bosch is a plant scientist with an interest in plant cell wall dynamics and plant reproduction. His cell wall-
related research focuses on the identification of the molecular and biological features underpinning cell wall recalci-
trance to sugar release, and elucidating how different environmental conditions affect cell wall-related traits relevant
for plant growth and development, biorefining, and ruminant digestion.
Prof. Iain Donnison is a plant scientist and the project lead for the Plants & Architecture NRN. His research interests
span the biology and breeding of crops, the sustainable use of natural resources, the matching of feedstocks to different
end uses, and the environmental impact of land-use change. He is also committed to working with industry to ensure
that academic research is translated into products and processes to deliver economic and environmental benefits.
Dr Morwenna Spear is a materials scientist with an interest in forestry, wood science and natural fibre composites. She
has worked on a wide range of projects in the bio-based materials field, ranging from wood modification by chemical,
thermal and polymer impregnation methods, to the development of tailored microstructures in natural fibre
reinforced polymer composites, controlling mechanical properties.
Dr Graham Ormondroyd is a materials scientist with an interest in BioBased Materials and a speciality in timber and
timber-based products. His research is focused on the use of bio-based materials as a replacement for synthetics in
various applications including construction and the valorisation of lignocellulosic waste materials.

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Appendix: A brief glossary

HYPOCOTYLS
The stem region of a seedling below the cotyledons (seed leaves).
MIDDLE LAMELLA
The thin layer that connects two plant cells and is rich in pectin.

MATRIX POLYSACCHARIDES
Complex polysaccharides found in the space between cellulose microfibrils. They are traditionally divided into pectins
and hemicelluloses.
POROSITY
Property that indicates how readily gases, liquids and other materials can penetrate an object.
PECTINS
Group of complex polysaccharides that are extracted from the cell wall by hot water, dilute acid or calcium chelators.
They include homogalacturonan, rhamnogalacturonans I and II, galactans, arabinans and other polysaccharides.
PRIMARY CELL WALL
The flexible extracellular matrix that is deposited while the cell is expanding.
SECONDARY CELL WALL
The flexible extracellular matrix that is deposited while the cell is still expanding is known as the primary cell wall.
When expansion ceases, a secondary wall is sometimes laid down inside the primary wall, making it stronger.
TURGOR PRESSURE
Force generated by water pushing outward on the plasma membrane and plant cell wall, that results in plant rigidity.
The loss of turgor pressure causes wilting.
102 Y. XING ET AL.

TRACHEARY ELEMENTS
Specialized cells in the xylem of vascular plants that are responsible for the conductance of water as well as providing
mechanical support.
VACUOLE
A membrane-bound cellular compartment, usually filled with a dilute watery solution. Mature plant cells often have
very large central vacuoles.
XYLEM
A tissue that comprises a group of specialized cells that are involved in the transport of water and solutes in vascular
plants. Mature xylem vessels essentially contain only the cell wall.