bearing materials in these regions (e.g., triangular supports in
corners of rectangular picture frames). A more efficient way to
stabilize a rigid framework is through triangulation such that
each strut is oriented so as to constrain the joint to a fixed
position. For these reasons, Nature repeatedly selected out
geodesic tensegrity systems. A more elegant and economical
solution to this problem that results in both structural efficiency
and flexibility is to impose an internal tension (prestress) to
reduce the play in the joints. Hence, Nature soon discovered
prestressed tensegrity structures as well (Table 1).
In summary, because it is the most economical solution to
the design challenges created by spatial constraints, energy
minimization rules, and the need to efficiently balance forces,
tensegrity is the way Nature builds. It is the only building
principle that can provide a single explanation (Occam's razor)
for how both unicellular organisms with geodesic exoskeletons
(e.g., radiolaria) and huge behemoths with prestressed
musculoskeletons (e.g., dinosaurs) could ever come into
existence.
Autocatalytic sets and solid-state biochemistry
Another concept that may be relevant to how life originated is
the idea is that once structures come about that can mediate
catalysis, then coherent self-reinforcing webs of chemical
reactions or ``autocatalytic sets'' will spontaneously em-
erge.
(18)
For example, if a primitive catalyst (protoenzyme) 1
accelerated the formation of protoenzyme 2 and 2 accelerated
the formation of 3, and so on, then at some point protoenzyme
X would emerge that could catalyze formation of protoenzyme
1. Loop closure would result in self-reinforcement of this
particular web of interactions. Thus, this particular cluster of
interacting components would increase in abundance relative
to molecules that were excluded from the web (Table 1).
On one hand, if autocatalytic sets were to form in solution
(as commonly assumed), then huge numbers of reactions may
be required before self-organization would be observed. On
the other hand, if some of these primitive protoenzymes
catalyzed the assembly of 3D scaffolds that bound the
interacting molecules and brought them in close proximity,
then the likelihood that the product of one reaction could
function as a substrate for a neighboring enzyme would
increase dramatically. Incorporation of this form of solid-phase
or ``solid-state'' biochemistry would therefore greatly increase
the probability that an autocatalytic set would develop and be
sustained over time (Table 1).
In fact, one of the major limitations of past theories of the
origin of life is that they failed to consider the importance of
solid-state biochemistry. The cytoskeleton of eukaryotic cells
orients most of the cell's organelles and many of the enzymes
and substrates that make up its metabolic machinery.
(14,19,20)
This increases the efficiency of chemical reactions because
substrate availability is not diffusion-limited in this context and
much larger molecules that would normally be insoluble or
spatially restricted by diffusion may be funneled into complex
metabolic pathways.
(21)
It also provides a mechanism to
integrate structure and function such that cells can respond
directly to environmental stresses (and scaffold deformation)
by altering cellular biochemistry
(22±24)
and strengthening
the scaffold against disruption.
(3)
Importantly, prokaryotic
cells also appear to use solid-state biochemistry on primitive
Table 1. Natural design principles
* Minimize energy expenditure. Nature is most economical and, thus, all natural structures tend to minimize energy and mass.
* Obey spatial constraints. Certain topologial rules constrain the possible forms that matter can take on, regardless of size or position.
* Develop emergent properties through architecture. At every size scale, complex properties and functions emerge from the behavior of the
ensemble; the material properties of any single element is much less important than how the different elements are joined and positioned in 3D.
* Establish a mechanical equilibrium. Architectural stability requires the establishment of a global balance of mechanical forces, although local
regions may be stressed.
* Use discrete networks. Nature does not use bulk solids to build; discrete porous networks offer greater structural efficiency and versatility.
* Maximize tensile materials. Disproportionate use of compression elements puts greater demands on energy (and eventually metabolism) for their
production, support and movement as the relative distance between interacting components increase.
* Stabilize through triangulation or prestress. Prestress and triangulation provide more efficient ways to stabilize discrete networks. Triangulation
results in stiff structures whereas prestress provides both flexibility and strength.
* Use structural hierarchies. Structural efficiency is maximized and evolution accelerated through the use of hierarchical networks, which are
themselves discrete structures on a smaller scale.
* Develop self-renewing functional webs through emergence of autocatalytic sets. Once molecules with catalytic functions appear, self-reinforcing
webs of chemical and structural interactions will spontaneously come into existence and expand at the expense of other non-interactive components.
* Enhance functional efficiency through solid-state biochemistry. Most of the chemical and enzymatic functions carried out by living systems
proceed on insoluble scaffolds using solid-phase catalysis. This increases the efficiency of chemical reactions, stabilizes functional networks, and
allows entire metabolic systems to self-assemble with others to create hierarchical structures with enhanced functionality.
Problems and paradigms
1162 BioEssays 22.12