• Behavioral differences may have genetic component • A part of brain (lateral frontal cortex) important in human intelligence • Positron emission tomography: a technology to visualize brain cell activity through measuring the extent of blood flow to particular brain parts • Differences in human behaviors (like intelligence, emotions, mood, anxiety levels) have been linked to genetic variations. Evolution and behavioral development Garter Snake In California, coastal and inland populations of garter snakes differ in diet. Coastal garter snakes feed on banana slugs. Inland populations eat other foods e.g. fish and frogs. • Is there a genetic basis to the behavioral differences in diet preference? Steve Arnold studied populations to see if there were genetic differences between them.
Brought pregnant females into lab and
reared babies in isolation.
Offered slug pieces to snakes.
Inland snakes usually ignored slugs, coastal snakes usually ate it. Then tested newborn snakes on response to odors.
Offered snakes flavored cotton swabs,
counted number of tongue flicks made. Coastal individuals responded much more strongly to slug odor. Arnold carried out heritability studies.
Found little variation within either population.
Preference for slugs within a population
largely fixed. Ultimate Explanation • Inland population don’t have slugs, but are exposed to leeches. – Active selection against leech-like /slug consumption. • Why? Cause leeches will eat a snake from the inside out! – There are few leeches in the coastal population, therefore no active selection against sluggy things. • What does the fact that some inland snakes will eat slugs mean to you about the populations? Crosses between populations produced individuals with wide variation in response to slugs.
Suggests differences between populations
have strong genetic component. What is evolutionary basis for difference Advantage to slug eating on coast is between populations? obvious.
Rare gene for eating slugs would spread
rapidly. Density affects the foraging behavior of fruit fly larvae • Larvae reared in lab for many generations under low density conditions move shorter distances in the same period of time than larvae from populations reared under high- density conditions (see Figure 3.23) Artificial selection and behavioral evolution • Testing proposition that behaviors evolve • Artificial selection experiments • Particular alleles responsible for phenotypes • Phenotypes allowed to reproduce…responsible alleles will become more common in experimental populations • While phenotypes linked to less successful alleles will decrease • So, evolutionary change can be seen in the laboratory through artificial selection experiments Nest building behavior in house mice • In natural conditions use grasses and other plant material for nest building • In lab conditions, happily accept cotton as a nesting material • The amount of cotton a mouse collects can be quantified as the number of grams it pulls into a nest cage over a 4-day period Artificial selection and behavioral evolution • Nest building in mice. • In starting population mice used 13-18 grams of cotton to line their nests. • To test whether this variation in building is caused by genes or environment • Carol Lynch selected for “high”, “low” and “control” groups of mice. • After 15 generations amounts of cotton used by mice were: – Controls 15g same as original population. – High 40g – Low 5g – In figure on next slide low, control, and high values are average for each generation – Conclusion: experimental populations evolved Other examples of Artificial Selection • William Cade has selectively bred crickets to sing rarely or almost continuously.
• In only two generations of artificial
selection Pulido et al. showed that the timing of migration in European Blackcaps could be delayed more than a week. Behavior is Subject to Selection
• In fact, genetic variation accounts for
about one-third to one-half of the variation in the onset of migratory activity in southern German blackcaps Artificial Selection • Artificial selection experiments show that (i) behavior is subject to selection and (ii) populations contain sufficient genetic variability to evolve rapidly. Adaptability of behavioral development • As we have seen the development of behavior such as singing can be strongly influenced by environmental effects. • It is important for development of behavior to be resilient to disturbance so that normal behavior can develop as often as possible. • Process of developmental homeostasis reduces effects of disturbance. Adaptability of behavioral development • Harlow’s “experiments” with Rhesus monkeys showed that rhesus monkeys entirely deprived of contact with mothers and “reared” by artificial surrogates gained weight and grew normally but behaviorally developed abnormally (BIG surprise!) and were terrified of other monkeys if exposed to them. Adaptability of behavioral development • However, monkeys given even 15 minutes contact a day with other young monkeys developed essentially normally. Adaptability of behavioral development • As adults, these monkeys interacted normally with others unlike those individuals which had no social contacts as infants, which were withdrawn or very aggressive. Developmental homeostasis in human embryonic development • Studies on songbirds show that adult ability to learn and sing complex songs is affected by food deprivation during period of rapid nestling growth.
• In humans food deprivation of the mother
during pregnancy appears not to have major effects on the fetus’ intelligence. Symmetry and attractiveness • Many organisms are influenced by how symmetrical other individuals are when making mate choices.
• Asymmetries are believed to be caused by
problems in development and symmetry thus signals an ability to overcome developmental challenges. Symmetry and attractiveness • There is some evidence that humans include symmetry in their ratings of an individuals’ attractiveness, but data are not conclusive and there is considerable debate. • Female brush-legged wolf spiders, however, are significantly more likely to mate with males whose foreleg hair tufts are symmetrical than those who are not. • Study was carried out using video images of spiders that were identical apart from the digitally manipulated foreleg tufts. • So, symmetrical body is an adaptive phenotype which increases reproductive success. However, in a number of species, two or three disticnt alternate phenotypes coexist comfortably. These arise as a result of the environmental differences among the individuals from a single species Polyphenism • In many species multiple alternative phenotypes occur in the same species (i.e. distinctly different body types occur).
• These different phenotypes arise as a
result of environmental effects from a single genotype. The environmental influence sends the developing organism down one or another distinct developmental pathway. • By which mechanism? Adaptive Value of Developmental Switch Mechanisms • Developmental switch mechanisms can produce polyphenism within same species. • Figure 3.32 • Different phenotypes can arise when dev sw mech are activated through in response to critical environmental cues. • See the figure for broadly variable phenotype, highly canalized phenotype, and polyphenism • Polyphenism may be a) food-induced (in castes of ants and other social insects b) socially-induced (in territorial and nonterritorial forms of cichlid fish) and c) predator- induced (in soldier caste of some aphids which possess more powerful grasping legs and a larger stabbing probiscis than nonsoldier forms) Developmental flexibility in Tiger Salamanders • A representative example of polyphnism is developmental flexibility in tiger salamamanders • Larvae live in ephemeral ponds.
• Most follow “normal” development and eat
small invertebrates. But some become cannibals (bigger with larger teeth) and eat smaller salamanders. Normal form Cannibal What factors affect decision to become a cannibal? • Ideas? What factors affect decision to become a cannibal?
• Relatedness of cannibal to others in pool.
• Density of salamanders • Size distribution of salamanders • A salamander surrounded by lots of non- kin can benefit by becoming a cannibal.
• By growing faster it can escape from the
pond sooner.
• Developmental flexibility allows
salamander to adjust its development if conditions are suitable, but not otherwise. Behavioral flexibility • Many animals choose among different behavioral phenotypes depending on environmental circumstances.
• For example, in many fish males adopt different
roles depending on their size and status. Large males defend territories but smaller satellite males act as sneakers darting in and releasing sperm whenever the dominant male mates with a female. Behavioral flexibility in Haplochromis burtoni • In cichlid fish Haplochromis burtoni. territorial males are brightly colored and satellite males are dull.
• Behavior is related to brain structure.
Behavioral flexibility expensive.
Should only evolve when benefit
outweighs cost. Satellite male
Territorial male Behavioral flexibility in Haplochromis burtoni
• GnRH (gonadotropin releasing hormone)
neurons in hypothalamus are 6-8 times larger in territorial males than in satellites.
• GnRH neurons stimulate testes
development and aggressive behavior. Behavioral flexibility in Haplochromis burtoni • Interestingly, GnRH size is variable.
• If male loses territory his neurons shrink
and he becomes dull colored.
• If territory opens up, male enlarges neurons,
switches to aggressive territorial mode • See figure 3.34 Behavioral flexibility in Haplochromis burtoni • Unpredictable social environment favors flexibility in Haplochromis.
• Neuronal flexibility allows males to adopt
best strategy for conditions. Adaptive Value of Learning • Learning is an adaptive modification of behavior based on experience • Its also a polyphenism because it too confers a highly focused behavioral flexibility that requires developmental modifications in the nervous system. • Selection favors learning when benefit outweighs the cost • Learning major element in behavioral flexibility. Ability to make use of experience in adjusting behavior can be selectively very advantageous. Learning in Australian Thynnine wasps • Males search for females based on pheromones they produce.
• Orchids possess flowers with a female decoy
petals that attract males using a similar scent.
• Orchid mimics female wasps appearance too.
When male lands and attempts to mate he gets a surprise and orchid gets a pollinator (pollen sacs stick to male’s back) Male tries to mate with “female wasp” on orchid, instead acquires a pollen sac.
Female wasp Fake wasp
Pollinia (poolen sac) on wasp’s back Wasps don’t learn to avoid orchids in general.
But, do learn to avoid orchids they have
visited before. Learning in Australian Thynnine wasps • What do wasps learn?
• Orchid’s location or orchid’s scent?
• Speculation: If wasps could learn scent
differences how plants benefit from having wasps revisit them? Costs and benefits of learning • For learning and flexible behavior to spread by natural selection, benefits must exceed costs.
• Benefits are the ability to exploit the
environment more effectively.
• What are the costs?
Costs and benefits of learning • Costs: Major cost is additional energy required to make/maintain neuronal tissue. Example of cost:
West coast marsh wrens sing more songs
(100) than east coast marsh wrens (40). Song system in brain weighs 25% more in west coast birds. Costs of large brain in humans • In humans: Brain 2% of body weight.
• But requires 15% of body’s oxygen and
20% of energy.
• Other costs to large brains in humans?
Costs of large brain in humans • Difficulty in giving birth. Behavioral flexibility expensive.
Should only evolve when benefit
outweighs cost. Clark’s Nutcracker seed-storing specialist. Much better at remembering where Something is located than other crows. Not better at remembering colors, a non- spatial task. Memory skills determined by bird’s needs. Intraspecific variability in learning • Behavioral flexibility does not only differ between species.
• In species where males and females
experience different selection pressures, differences in learning ability occur. Spatial learning in voles • Male meadow voles are polygynous. Their home ranges are 4X larger than those of females.
• Prairie voles are monogamous. Males
and females have same size home ranges Spatial learning in voles • Spatial learning abilities of voles was tested in mazes.
• Male meadow voles made significantly
fewer mistakes than female meadow voles, but there was no difference between the sexes in the performance of male and female prairie voles. Spatial learning in voles • Male meadow voles have a large hippocampus than females and this area in the brain processes spatial information.
• However, the enlarged hippocampus only
develops during the breeding season. Spatial learning in cowbirds • Female cowbirds parasitize other birds nests. They need to be able to remember nest locations and monitor them over time. Female cowbirds have a larger hippocampus than males. No difference between sexes in related, but non- parasitic grackles and red-winged blackbirds. Operant conditioning • Spatial learning not only behavior that has clearly been shaped by selection.
• In operant conditioning (or trial and error
learning) an animal learns to associate an action with its consequences.
• E.g. a rat pushes a lever and gets a food
pellet. Operant conditioning • Usually a behavior and its consequences must be closely linked in time for conditioning to occur.
• However, in rats tasting novel foods
nausea that occurs several hours after eating a food will be associated with that food, which in future will be avoided. Another example of operant conditioning is that predators learn to recognize noxious prey after tasting them.
How does warning
coloration benefit animal that is tasted if it is killed? Does operant conditioning occur in humans?