One challenge for
the theory of evolution: it needs to encompass both creativity and
predictivity. Creativity, because new species come into being, never having
been observed before. Predictivity, because underlying causal mechanisms come
into play as organisms with certain traits are selected for instead of their
less adaptable relatives. In addition, biological arguments often invoke the
notion of ‘function’ – which is an overlying rather than an underlying
cause hierarchically. Biochemical arguments, excising macroscopic ecology from
the picture, attempt to explain the function in terms of ‘structure’ to
emphasize underlying causes.
In his treatise,
Jeffrey Wicken incorporates thermodynamics and information to broaden the
theory of evolution beyond the neo-Darwinian. Chapter 12 of his book is
titled “The Economics of Selection”. How do organisms grow and reproduce? By
funneling energy through themselves and degrading it to build new structures.
Why do they so? Well, to grow and reproduce. Note the chicken-and-egg argument I
just made with the how and why questions, which has led to the idea of
biological relativity – no particular level within the hierarchy can be easily
singled out as the primary cause.
The economic
currency here is energy. In our neighborhood of the solar system, an energy
gradient exists between the sun (shooting out photons) and the ‘coldness’ of the
vacuum of space. In between sit the planets, full of matter and some (okay,
one) with the potential for supporting living organisms. Translating the energy
gradient in chemical terms, Wicken writes: “As energy penetration increased the
free energy of the emerging biosphere, it built into it proximate free energy
gradients that could be exploitatively tapped. These involve both sources with
high-energy electrons and oxidative sinks for accepting those electrons. The
fundamental economic boundaries of a system are set by its source-sink
relationships and the kinetic mechanisms at its disposal for exploiting them.”
My chemistry
students will hopefully recognize that some sort of redox reaction must be
involved. Electrons are moving from a source (the reducing agent) to the sink
(the oxidizing agent). Hydrogen is the prototypical reducing agent. Oxygen is
the prototypical oxidizing agent. Using handwaving (or bookkeeping) arguments
involving ‘oxidation numbers’, students learn in introductory chemistry that
hydrogen transfers its electron to oxygen and H2O is formed from
elemental hydrogen and oxygen. The formation of water is energetically very
favorable – there’s a gradient from the higher (potential) energy reactants, H2
and O2, to the lower (potential) energy product H2O. More
subtly, and I emphasize this point in my upper division P-Chem thermodynamics class,
O2 is the “high-energy” species from the perspective of the relative
‘strengths’ of chemical bonds. And why is carbon central to life? I’d
say it’s along for the ride thanks to its ability to expand chemical space to
create new structures. This notion of expanding chemical space will be
important to Wicken’s thermodynamic argument.
While our present atmosphere
has lots of free O2, and heterotrophic organisms (we can’t make our own
food, we have to eat it!) utilize O2 for metabolism, this was not so
at life’s origin on our planet. Oxygen was locked into both minerals (many ores
are metal oxides) and carbon (as carbon dioxide in the gas phase, bicarbonate
in aqueous solution, and carbonate both dissolved and in ionic solids). The
reason why we have lots of O2 today is because, at some point in the
distant past, photosynthesis was ‘invented’ thereby allowing autotrophic organisms
(that make their own food) to directly utilize the sun’s high-energy photons to
reduce CO2 essentially by adding hydrogen to it. That’s why living
organisms are full of compounds containing carbon, hydrogen, and oxygen.
Carbohydrates and lipids are in this category. (Proteins and nucleic acids also
contain some nitrogen, sulfur and phosphorus.)
As pathways to utilize
energy emerge, there will be competition. This is where thermoeconomics comes in.
Wicken writes: “Competition among thermodynamic flow patterns expresses itself
in ecosystem dynamics by imposing a condition of selection on community
structures that in turn imposes selective conditions on adaptive strategies of
populations. No community organization is uniquely able to process the energy
made available by a given abiotic matrix. Any community is but one of many
alternative solutions to the business of energy processing, and is therefore in
competition with these alternative solutions.”
And how do we
expect evolution to proceed in this case? Here’s what Wicken has to say: “Since
the biosphere is a closed system, and since irreversible flows of energy
through closed systems require cyclic movements of matter… ultimately all
[autocatalytic pathways] must contribute to the cyclic movement of matter
through the biosphere. As open systems, organisms fulfill their thermodynamic destinies
by fitting into higher-order systems that express greater degrees of closure.
The ecosystem, with its community structure for processing energy, is the first
level of cyclic closure – cycling scarce nutrients and unidirectionally
processing others, such as carbon and oxygen, which are in greater supply. Then
come systems of ecosystems, in which the outflow of one is utilized as input by
another. The biosphere as a whole is the ultimate unit of cyclic closure.”
This makes sense
to me as a physical chemist. Using the language of thermodynamics allows us to
deal with macroscopic-level variables without being bogged down by microscopic
causal explanations. We just need to follow the energy trail! Wicken’s distinguishing
the fates of C and O (and implicitly H, the most abundant element in the
universe) from N, S, P, and trace metals is, I find, a refreshing way to look
at the interplay of the different elements in chemistry. That’s why we have a
nitrogen cycle! Cycles are fundamental to life.
But there’s more.
Wicken writes: “Yet ecosystems are quite vulnerable thermodynamically to
invasive dissipative modes – whether nurtured within the relational structure
of the community or by endogenously generated parasites.” There’s a reason why
life is dynamic: the second law of thermodynamics. Entropy must continue to
increase in the biosphere (approximated as a closed thermodynamic system) if it
is not stuck in chemical equilibrium ‘death’. This thermodynamic vulnerability
of an ecosystem is a feature, not a bug, in the evolution of life. I expect this
principle of vulnerability to be dominant at life’s origins, before the
establishment of complex control mechanisms in organisms to stay alive.
As an ecosystem
matures, the flows change. Wicken argues: “That mature ecosystems have lower
specific dissipations [i.e., lower specific entropy production] than immature
ones (higher biomass/throughput ratios) reflects the selective premium on using
resources efficiently. That evolution occurs under conditions of limited
resources means that it occurs under economic boundaries with limiting kinetic
means of degrading energy.” What this means is that in the early immature
stage, there tend to be multiple kinetically accessible pathways to dissipate
energy. New pathways (the “invasive modes”) continue to emerge, essentially to
expand the energy economy. But gobbling up resources leads to shortages, leading
to the selection of more efficient routes – and the closing down of some
pathways. This interplay leads to specialization and mutualism. Wicken writes: “In
ecosystem development, founder populations of energy-intensive generalists give
way to more complexly organized communities of interacting specialist
populations that get more biomass for their energy dollar.” This also applies
to a complex multicellular organism such as humans! Wicken argues that our
natural life span corresponds to the decreasing specific entropy production from
early stages of the zygote to eventual death of the organism.
Wicken subdivides
this into three stages: (1) “In early development, an organism’s high
dissipative rate and low efficiency provide thermodynamic force for the
generation of organization according to its coded information.” (2) “Maturation
is accompanied by decreasing specific dissipation as body size peaks and
organizational networks reach their full, mutualistic complexification.” (3) “The
decline of the dissipation during the aging process expresses a gradual
yielding of homeostatic function… systematic losses of the behavioral and
metabolic plasticity that allow organisms to accommodate environmental stresses.”
But why do organisms
eventually lose homeostatic ability? Why can’t we stay immortal keeping
ourselves at steady state and away from equilibrium? Why is death built into
life? Wicken writes: “The wearout of energy-processing pathways inheres to organisms
by virtue of their particular condition of closure, which is to information
rather than energy.” We’ll need to unpack this statement. First, the environment
surrounding an organism is ever-changing. To maintain itself, an organism needs
to buffer itself against many of these changes. An energy-inefficient
generalist (with lower information content) has a variety of mechanisms it can
employ to respond to external perturbations. An energy-efficient specialist within
a mutualist community (with higher information content) has fewer response
mechanisms. Don’t forget that specializing and mutualism is to some extent
inevitable because of the second law. Wicken writes: “Since success in the
competition for energy flows is predicated on current adaptive payoff
rather than the future stabilization of community structures, the
survival-reproductive fruits of minimizing metabolic and behavioral burdens
make the trend toward increasingly specialization inexorable… This essential
conflict between adaptive commitment and environmental change mandates eventual
extinction for most species.”
So on the one
hand, thermodynamics drives the opening up of chemical space and encourages a
diversity of structures to form. The more different kinds or types, the higher
the entropy produced. And entropy must be produced for anything to proceed.
Otherwise you’re stuck in chemical equilibrium – the stasis of death (or I
suppose an unchanging immortality). You’re down in the (potential energy) well
and there’s nowhere to go. But as you open up the chemical space and increase
in complexity, specialization and mutualism kick in for increased efficiency in
a climate of ever-changing resources. But specialization is also an eventual
death-knell in a dynamic environment. All those command-and-control systems
that emerged allowing an organism to grow (and possibly reproduce) become
limiting because they are selected here-and-now and not for the future. We’re
all short-termers at heart. To build robustness for the long-term requires an
efficiency trade-off, and the possibility of losing the fight for resources in
the here-and-now. A Catch-22.
Why are we in this
situation? Here’s Wicken’s overaching view: “The driving force [for this
evolutionary trend, in both the organic and socioeconomic realms, has been the
fact that energy resources have existed in excess of their mechanisms of
utilization. Evolution has been more kinetically than energetically limited.
Prior to the emergence of technology, this kinetic limitation was guaranteed.
Relatively small fractions of influx in solar radiation are autotrophically
fixed. Of that fraction, not all can be heterotrophically utilized – the residue
sinking into fossil fuel. Socioeconomic evolution has powerfully accentuated
the trend toward the invention of kinetic mechanisms to create new patterns of
dissipation. Econodynamics is inescapably evolutionary in this way. A caveat
is that these dynamics have promoted the evolution of more highly dissipative
economies by exploiting the fossil fuels that resulted from eons of surplus
production of autotrophic fixation. As future socioeconomic evolution becomes
more energy-limited, selection for efficiencies will inevitably reverse the
historical relationship between organizational complexity and dissipation.”
Humans – we’ve really changed things on Planet Earth!
I close this long
meandering blog with closing thoughts by Wicken on the relationship between
parts and the whole: “The whole is not exactly more than the sum of its parts,
since parts are relationally constituted by the wholes in which they
evolved… The organic world consists of just such mutually constituting
part-whole relationships between individuals and higher-order flow patterns.
The success of any whole is predicated on the coordination of its parts;
conversely, adaptive strategies of parts are conditioned by the requirement
that they participate in higher-order webs of processing energy. The
information content of an ecosystem expresses this hierarchically codefining
relationship between part and whole.”