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Dissertation Proposal - The University of Arizona Campus Repository

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ABSTRACT<br />

<strong>The</strong> fitness <strong>of</strong> any evolutionary unit can be understood in terms <strong>of</strong> its two basic<br />

components: fecundity and viability. <strong>The</strong> trade-<strong>of</strong>fs between these fitness components<br />

drive the evolution <strong>of</strong> a variety <strong>of</strong> life-history traits in extant multicellular lineages. Here,<br />

I show evidence that the evolution <strong>of</strong> germ-soma separation and the emergence <strong>of</strong><br />

individuality at a higher level during the unicellular-multicellular transition are also<br />

consequences <strong>of</strong> these trade-<strong>of</strong>fs. <strong>The</strong> transition from unicellular to larger multicellular<br />

organisms has benefits, costs, and requirements. I argue that germ-soma separation<br />

evolved as a means to counteract the increasing costs and requirements <strong>of</strong> larger<br />

multicellular colonies. Volvocalean green algae are uniquely suited for studying this<br />

transition since they range from unicells to undifferentiated colonies, to multicellular<br />

individuals with complete germ-soma separation. In these flagellated organisms, the<br />

increase in cell specialization observed as colony size increases can be explained in terms<br />

<strong>of</strong> increased requirements for self-propulsion and to avoid sinking. <strong>The</strong> collective<br />

flagellar beating also serves to enhance molecular transport <strong>of</strong> nutrients and wastes.<br />

Standard hydrodynamic measurements and concepts are used to analyze motility (self-<br />

propulsion) and its consequences for different degrees <strong>of</strong> cell specialization in the<br />

Volvocales as colony size increases. This approach is used to calculate the physical<br />

hydrodynamic limits on motility to the spheroid colony design. To test the importance <strong>of</strong><br />

collective flagellar beating on nutrient uptake, the effect <strong>of</strong> advective dynamics on the<br />

productivity <strong>of</strong> large colonies is quantified. I conclude first, that when colony size<br />

exceeds a threshold, a specialized and sterile soma must evolve, and the somatic to<br />

8

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