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The Origin and Evolution of Mammals - Moodle

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elevation <strong>of</strong> maximum aerobic activity. Virtually all<br />

authors have made the assumption that such a<br />

complex arrangement could not have evolved in its<br />

entirety in one step, <strong>and</strong> therefore must have arisen<br />

by a simpler route consisting <strong>of</strong> two successive<br />

steps. <strong>The</strong> argument then shifts to which function<br />

was initially selected for, <strong>and</strong> which only subsequently.<br />

<strong>The</strong> underlying assumption is arguably<br />

false, however, <strong>and</strong> therefore the ensuing question<br />

is the wrong question. <strong>The</strong> correct question is not a<br />

matter <strong>of</strong> the order in which the parts <strong>of</strong> the complex<br />

whole changed, but rather <strong>of</strong> how the various<br />

evolving characteristics are interrelated such that<br />

every stage in the transition from fully ectothermic<br />

organism to fully endothermic organism remained<br />

a viable, integrated entity. This approach to<br />

underst<strong>and</strong>ing the origin <strong>of</strong> mammals is taken<br />

up a fortiori later, but for the moment the particular<br />

case <strong>of</strong> the origin <strong>of</strong> endothermy can be illustrated<br />

by way <strong>of</strong> a plausible scenario (Fig. 4.13). Suppose<br />

a mutation occurred that caused a small increase in<br />

number <strong>of</strong> mitochondria in all the cells. This would<br />

increase the maximum level <strong>of</strong> sustainable aerobic<br />

activity a trifle, permitting a few extra minutes in<br />

the chase. Simultaneously <strong>and</strong> unavoidably, the<br />

greater heat produced by the extra mitochondria<br />

would incrementally increase the animal’s average<br />

body temperature by a few degrees, perhaps enough<br />

to permit an extra half an hour’s activity before torpor<br />

set in at nightfall. Not much else can happen<br />

now until maybe a small incremental increase in<br />

oxygen uptake, sufficient to support another increment<br />

in BMR, or perhaps a modification to the skin<br />

vascularisation reducing its conductivity slightly.<br />

A hypothetical model like this indicates that both<br />

the main functions <strong>of</strong> endothermy can in principle<br />

evolve simultaneously <strong>and</strong> incrementally, with neither<br />

having primacy over the other. Furthermore,<br />

within the constraints imposed by the functional<br />

integration <strong>of</strong> organisms, it is actually a much simpler<br />

explanation than that based on the serial accumulation<br />

<strong>of</strong> separate functions.<br />

Meanwhile, in this light the question <strong>of</strong> when<br />

endothermy evolved becomes transmuted into the<br />

question <strong>of</strong> what level <strong>of</strong> endothermy had been<br />

achieved by this or that particular stage. <strong>The</strong>re is little<br />

doubt that the pelycosaur-grade synapsids were<br />

ectothermic, as witness particularly the dorsal sail<br />

EVOLUTION OF MAMMALIAN BIOLOGY 129<br />

<strong>of</strong> Dimetrodon <strong>and</strong> Edaphosaurus. At the other<br />

extreme, a case has already been made that cynodonts<br />

had achieved a relatively high level <strong>of</strong><br />

endothermy. <strong>The</strong> largest problem is assessing the<br />

status <strong>of</strong> pre-cynodont therapsids such as gorgonopsians,<br />

therocephalians, <strong>and</strong> dicynodonts. <strong>The</strong><br />

impressive modification <strong>of</strong> the locomotory mechanics<br />

at the basal therapsids level supports the view<br />

that there were enhanced aerobic activity levels.<br />

<strong>The</strong>ir widespread abundance in temperate regions<br />

<strong>of</strong> the world suggests a reasonable degree <strong>of</strong><br />

thermoregulation.<br />

As to the temperature physiology <strong>of</strong> the earliest<br />

mammals themselves, the story is complicated by the<br />

process <strong>of</strong> miniaturisation that occurred in the lineage<br />

which led to them. Very small organisms face an<br />

array <strong>of</strong> biological constraints <strong>and</strong> potentials differing<br />

in several respects from those <strong>of</strong> otherwise comparable<br />

larger ones, as will be discussed presently<br />

(page 135). Nevertheless, it is hard to doubt that animals<br />

with the complete anatomy <strong>of</strong> modern mammals<br />

did not share their complete general physiology<br />

as well.<br />

An integrated view <strong>of</strong> the origin <strong>of</strong><br />

mammalian biology<br />

Having reviewed what can be inferred from the<br />

fossil record about the evolution <strong>of</strong> the mammalian<br />

condition <strong>of</strong> various separate functional systems,<br />

it is now appropriate to take an integrated view<br />

<strong>of</strong> the evolution <strong>of</strong> mammalian biology as a whole.<br />

Comparison <strong>of</strong> a typical modern reptile such as a<br />

lizard or crocodile with a primitive living mammal<br />

reveals immediately just how great was the transition<br />

from their last common ancestor to the first<br />

mammal, <strong>and</strong> how it affected virtually every physiological<br />

<strong>and</strong> anatomical feature. <strong>The</strong> evolutionary<br />

paradox implied by this observation has been long<br />

<strong>and</strong> widely appreciated; indeed, ever since evolution<br />

by natural selection became accepted as the<br />

overarching explanation for the diversity <strong>of</strong> life. On<br />

the one h<strong>and</strong>, evolutionary change is caused by<br />

mutations in genes that affect discrete features, yet<br />

on the other h<strong>and</strong> organisms must remain complex,<br />

tightly integrated individuals in which all the structures<br />

<strong>and</strong> processes are designed to interact with<br />

one another to generate the overall biological nature

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