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1 Chapter 10 Adaptation: from genes to traits p Cascades of Genes (p. 304) 1. Complex adaptations A. Coexpressed traits selected for a common function, 2. often a novel trait A. not inherited from an ancestor B. no obvious counterparts (homologs) in related lineages. C. E.g., ability to digest citrate, the eye, venom delivery systems, etc. 1 Complex adaptations (p. 304) 1. can require multiple mutations 2. Multiple components must be expressed together for the trait to function. ( irreducible complexity ) 3. existing genes expressed in a new developmental context 2 Regulatory Networks 1. Systems of interacting genes, transcription factors, promoters, RNA, and other molecules. (Fig. 10.2) 2. biological circuits controling the activation of genes 3. Hox genes encode transcription factors that determine the identity of body parts along a head to tail axis (Fig. 10.3) 4. Hox mutations can have major effects 3 1

2 10.2 Generating Innovations p Mutations can reprogram conditions in which a gene is expressed 2. Old proteins take on new functions without decreasing original function A. Promiscuous proteins, Paralogs, Gene Recruitment, Horizontal gene transfer 4 1. Promiscuous proteins carry out more than one function, A. mutations can improve the function of the protein for different tasks. 2. Paralogs A. Homologous genes that arise by gene duplication. a. Duplication can lead to new functions B. Unequal crossing over a. Globin gene family 5 Gene duplication can produce novel functions (Fig. 10.5) Promiscuous proteins are especially likely to take on new functions after duplication 2

3 Recruitment or co-option 1. Novel traits arise when existing genes are expressed in new developmental contexts 2. Mutations of regulatory genes can cause genes to be expressed in different tissues than before. Our activities have altered the natural environment p Sphingobium, a soil bacterium, has assembled a new metabolic pathway that can mineralize PCP (pentachlorophenol) by patching together promiscuous enzymes recruited from pre-existing pathways (Shelley Copley) 8 Evolution of novel traits Around generation 33,000, one population acquired the ability to metabolize citrate, a second nutrient in their culture medium that E. coli normally cannot use. (Fig. 10.6) 9 3

4 Venom Evolution: Borrowing Genes for Deadly New Jobs p Crotamine is a toxin present in the venom of rattlesnakes A. duplication and co-option of proteins originally involved with other functions. B. Beta defensins are antimicrobial peptides (Fig. 10.8) C. Gene duplication and co-option and snake venoms (Fig. 10.9) 2. Venom genes have been recruited from genes expressed in many organs of snakes (Fig ) 3. Venom evolved before snakes themselves (Fig ) The Genetic Toolkit p Evolutionary Developmental Biology examines regulatory genes that direct development 2. Hox gene products are transcription factors that bind to DNA and control the transcription of other genes. 3. Development in flies and mice is controlled by homologous Hox genes that evolved in a common ancestor > 570 million years ago (Fig ) 11 Dorsal ventral-patterning is conserved (Fig ) 1. vertebrate body plan is a dorsoventral inversion of the insect body plan. (Fig ) 4

5 1. Orthologs A. Homologous genes separated by a speciation event 2. Fly legs and mouse legs patterned by same genetic cascade (Fig ) 3. Expression differences in a single gene give rise to hind limb elongation in crickets and forelimb elongation in bats (Fig ) 13 Traits can disappear when the expression of a patterning pathway is altered, blocked, or even just interrupted. 1. Disruptions in Shh (Sonic hedgehog) expression cause failure of limbs to develop (Fig ) 2. Changes in limb patterning pathway result in limb loss in snakes (Fig ) A. HOXC8 and HOXC6 domains are expanded along the body axis in python embryos lengthening the trunk at the expense of the limbs 14 Complex eyes have evolved in several lineages (Fig ) 1. Dozens of different kinds of animal eyes. 2. Opsins are light sensitive proteins on the surface of photoreceptor cells 3. Opsins belong to a family of proteins called G-protein coupled receptors (GPCRs), also known as serpentine proteins. 4. Serpentine proteins transduce signals in the cells of eukaryotes 5. Opsin originated by duplication of a serpentine protein gene (Fig ) 5

6 Opsins evolved approximately one billion years ago (Fig ) 1. Crystallins evolved through gene recruitment (Fig ) A. proteins that form transparent lenses to focus light B. Recruited from ancestral genes that coded for heat-shock proteins C. a heat-stressed cell can be protected by being stabilized by small heat-shock proteins 2. the lens itself has heat-shock protein properties 3. Eyes may have evolved from simple patches of light-sensing cells on the skin to the complex structures we see today (Fig ) 10.9 Constraints on adaptation 1. Laws of physics 2. Pleiotropy A. Single gene affects expression of many traits 3. Antagonistic pleiotropy A. multiple effects of a gene have opposing effects on fitness B. Antagonistic pleiotropy: restricts the number of cervical vertebrae in mammals to seven (Fig ) 6

7 Complex adaptations are not perfect 1. Recurrent laryngeal nerve in mammals comes from the brain, through the neck, to the heart, round the dorsal aorta and up to the larynx, through the neck. 2. Nerve may be up to 4m long, despite optimal route of several inches. 3. Indirect route is result of evolution of mammals from fish, which had no neck and a short nerve that innervated one gill slit and passed near the gill arch. 4. The gill it innervated became the larynx and the blood vessel became the dorsal aorta. (Fig ) Convergent evolution 1. Independent evolution of similar traits in different lineages A. Result of similar selection pressures B. The process by which similar traits evolve in unrelated lineages. C. Convergent evolution in mammals 1. Parallelism A. Convergent evolution that arises through mutation of the same genes in different lineages 2. Deep homology A. traits in different lineages arise from same inherited regulatory networks B. These networks are often shared by many species and are modified to produce variations of the same trait across species, such as the eye, for example. C. The Pax 6 gene which underlies all eyes is found in dinoflagellates and mice and fruit flies 7

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