Midterm Flashcards
Evolution Before Charles Darwin
Enlightenment - no divine right to rule, natural phenomena are mechanically caused by external forces
Erasmus Darwin - Charles’ grandfather, organisms’ sensitivity to the environment shapes their minds and bodies, better forms have increased over time
Jean-Baptiste Lamarck
Jean-Baptiste Lamarck
- animals have an internal drive to physically accommodate to environmental conditions and become more complex
- persistently successful accommodations can become so habitual that they become physiologically based and get passed on to offspring
3 Influences of Darwin’s Theory
Charles Lyell (geologist) Thomas Malthus (economist/demographer) Alfred Russell Wallace (naturalist)
Charles Lyell
geologist, friend of Darwin
argued for a very slow and incremental pace of geological change
Thomas Malthus
economist and demographer
emphasized danger of uncontrolled human reproduction with limited resources
helped Darwin see the role of resource competition
Alfred Russell Wallace
independently discovered the theory of natural selection; letter prompted Darwin to submit their theories together in 1859
“Survival of the Fittest”
“fitness” is really “fittedness” - term was coined by Herbert Spencer, not Darwin
fitness is not determined by health or strength, but by reproductive success and ability to adapt to environment
3 dimensions of biological analysis
length - longitudinal analysis (homology)
breadth - comparative analysis (analogy)
depth - functional analysis (physiology)
How nature “backs up” her data
- Things naturally tend to fall apart and complicated mechanisms tend to wear out and fail
- One defense is to make copies
- Copying is never perfect - variation is inevitable
- Copying takes time and energy
- Some copying processes will be more profligate or successful than others
- Favored copying processes will be ones that best “fit” the conditions
Herbert Spencer
coined the phrase “survival of the fittest”
argued that evolution argued by Lamarckian use-inheritance
believed that learned habits could be passed on by reproduction –> social progress could lead to biological progress
argued that “primitive” societies were less evolved “races”
Haeckel and Progress
argued that evolution tends toward greater complexity
humans were presumed to occupy the highest rung on the biological ladder and were “the most evolved”
Why Evolutionary Misconceptions are Wrong
Evolution is not linear and not necessarily progressive.
Its tendency is to diversify and expand into novel niches.
Evolution spontaneously tends toward increasing complexity because there is nowhere else to go.
“Planet of the Apes” Fallacy
nothing about natural selection produces inevitable trends, even improvement
humans didn’t replace or improve upon apes
Types of Selection
stabilizing selection - selection against extreme phenotypes that deviate from the mean
directional selection - selection favors extremes
disruptive selection - either extreme is favored
eg Japanese pheasants
The Red Queen analogy
the environment is always a moving target
it can look like there is an “arms race” between organisms adapting to the same environment, but they are just trying to keep up with the same changing environment
sometimes you have to run fast just to stay where you are
Mendel’s Basic Discovery
simple traits are controlled by an interaction between the two “doses” of each gene type (allele) that are present in each plant
because of this, they can interact or mask each other’s effects (dominant/recessive)
independently assort in subsequent generation according to simple binomial rule
Protein Functions
catalysts - aid chemical reactions, eg digestive enzymes
structural elements - or link other molecules, eg skin and hair
recognition or signaling - for cell cell communication
molecular “chaperones” - for other molecules that bind or transport eg hormone receptors and hemoglobin oxygen transport
SNPs
single nucleotide polymorphisms; different people can have different bases in a specific location
can be used for identification, diagnosis, and tracing ancestry
Genetic “drift”
statistical sampling effects producing random selection
there is always a 50% loss in genes from each parent
sex leads to inevitable gene elimination by chance
Founder Effect
a small group initially founds a population and diversifies from there
reduces genetic diversity
Why shift to a genetic perspective?
all bodies eventually perish! what persists are the genes that contribute to those traits
but: the particular combination of genes and traits that constitute an individual will likely never recur, except for in identical twins and asexual species
eg. Secretariat effect
Sickle Cell Anemia
sickle cell is only produced by the genetic mutation if both hemoglobin A molecules are sick type
hetereozygotes have 50% hemoglobin beta exhibiting mutation, which does not cause anemia but a low level of cell damage
this is sufficient to expose the malaria parasite to the immune system and help eliminate it
so: sickle cell heterozygotes are positively selected for
balanced polymorphism
Mitochondrial DNA
not contained within the cell nucleus but within the organelle
mitochondria are passed on in the cytoplasm of the ovum and therefore come only from the mother
mtDNA has bacterial origin!
(Y chromosome DNA is similar - only passed through male lineage)
Y chromosome inheritance
long arm of the Y chromosome does not recombine in sexual reproduction
inherited intact from father to son
therefore transmitted in all or none fashion and can be traced from individual to individual
gene duplication –> functional divergence
eg hemoglobin:
spontaneous degradation of duplicate genes produced beta-hemoglobin variants with different oxygen affinities
problem of transferring oxygen from maternal to fetal hemoglobin provides “functional niches” that variant hemoglobins can fit
Regulatory Genes
produce proteins that control patterns of gene expression in the different cells of a body
Extended Phenotype
selectively favored effects are expressed outside the body that produces them
genes can produce molecules whose primary focus is to control another species’ development or behavior
Death as a Reproductive Strategy
trade-off point between maintaining a body and instead putting that energy into reproduction is different for different species/life stages
salmon kill themselves in an effort to arrive upstream ahead of competitors and produce hundreds of offspring
mantis males easier to be caught and eaten late in the breeding season when chances of reproducing are slim and nutrition from eating their bodies could aid female reproduction
Sex has evolutionary costs
- only a random half of one’s genome gets passed to each offspring, and some genes don’t get passed on
- good gene combinations tend to get broken-up in each generation (Secretariat effect)
- time and resources are required to locate a mate, and often in fighting for mates
- probability of encountering a potential mate is reduced by 50%
- males use up resources and don’t contribute to offspring nourishment and health
Sex requires social interaction
- over the course of evolution, sexual reproduction has played the dominant role in promoting social behavior and cooperative organization within a species
- by distributing mutually exclusive reproductive adaptations among different individuals the advantages and disadvantages of each are separated
- traded off against the costs of making autonomous reproduction impossible, resulting in different combinations of sexual traits
sexual reproduction involves a mix of:
- Recombination (genes)
- Anisogamy (sperm and ova)
- Gender (male and female)
Why recombination?
genes must work in combination with one another
necessary to have them linked to keep working combinations together during reproduction
but this risks having a damaged gene ruin the reproductive chances of any linked gene
periodic recombination (shuffling) allows genes to escape linkage with “bad” genes
Anisogamy
“not the same gametes”
different gamete phenotypes: usually haploid, as in ova and sperm
sperm selected for: high numbers, low production costs, easy dispersal, high genetic diversity –> small, mobile, expendable
ova selected for: robustness, ample metabolic resources –> large, immobile, easily protected
Hermaphroditism
(both kinds of gametes) requires a compromise with respect to gender-distinct bodies where both dispersal-variation and investment-protection can be independently maximized
usually in conditions where access to mates is reduced
Sex vs. Body Gender
sex - typically understood in terms of male and female, used to refer to either whole animal bodies or to the organs of reproductions in plants that carry sperm (pollen) or ova (egg cells)
gender - linguistic designation of nouns and pronouns typically into masculine, feminine, and neuter grammatical types; has come to be used by analogical extension to refer to masculine vs feminine “roles” aka GENDER IS A SOCIAL CONSTRUCT
Conditions that Favor Asexual Reproduction
minimal competition for resource
predictable conditions
higher reproductive rate, lower variation
Conditions that Favor Sexual Reproduction
significant competition for resources
unpredictable conditions
lower reproductive rate, higher variation
Blue-Headed Wrasse
sex change
all begin life as females
live in a school defended by a single male, who is much larger on average, has a distinct blue head, and inseminates all eggs of the school
if this male dies, the larger females will compete and one will transform into a male developing a blue head
Anemonefish
“clownfish”
have an ability to prevent anemones from stinging them
pairs “nest” in an anemone where they raise young who can feed off prey paralyzed by anemone
begin life as males capable of producing sperm
young males pair up with older females who have successfully defended an anemone “nest site” in which to raise the young
older female dies, younger male undergoes a sex change to become a female and gonads begin to produce ova
young males will be attracted to mate with female
Darwin’s Theory of Sexual Selection
explains the evolution of apparently nonfunctional, superfluous, or harmful traits including human intelligence
emphasizes Darwin’s focus on reproduction
basic logic - reproductive competition not only dissociates members of a species into distinct sexes, but can lead to competition within the same sex for access to reproduction
Darwinian Logic of Sexual Selection
- better reproduction is more important than optimizing survival or access to resources
- there are often major differences in an individual’s reproductive output
- possibility of increasing reproduction could lead to competition for mates
- traits that trade health and survival for reproductive advantages will evolve where not blocked by natural selection
Modern Logic of Sexual Selection
- sex that contributes more time, energy, and risk in support of offspring is “limiting resource” for other sex
- sex that invests more evolves greater discernment of mates; sex that invests less evolves a means to bias the selection in its favor by increasing the total number of mates
- manipulation of mating changes may occur via aggressive exclusion of competitors, competition over valued resources, promiscuity, risky displays that signal health or vitality
Displays as Handicaps
eg widowbirds
widow bird males have long tails that attract both females and predators
experimental manipulation shows that females prefer longer than normal tails
Tournament Reproductive Strategy
about 98% of mammals, sexual dimorphism, male energy largely devoted to competition and no male parental investment, high variance in male reproductive success
male display; female choice
high polygyny - rival males are excluded
Pair Bonding Reproductive Strategy
95% of bird species
sexual monomorphism, male energy divided between competition and parental investment, low variance in male reproductive success
mutual courtship
low degree of polygyny, some polyandry, mostly mated pairs
Male Defense: either resources or females
resource defense - males compete for territories that attract females, who spend more time there and are available for mating
female defense - food is widely distributed and females feed in groups, so males compete to prevent other males from mating with them
Bowerbirds
competing for mates using artifacts:
male bowerbirds attract mates with elaborately constructed and decorated displays (bowers) that only superficially resemble nests
females will mate in the bowers and then raise young on their own
males compete for attractive objects to decorate their bowers, often stealing from other bowers
Factors contributing to pair bonding
- offspring that require extensive or prolonged care to reach a stage where they can forage for themselves
- non-abundant, non-clumped food resources
- limited nest or burrow sites, or sites requiring defense or significant effort to produce
- low population densities
Bird and Mammal Cooperative Care
- cooperative rearing strategies are generally rare, but can be found in environments where resources (like nesting sites) are rare
- because defending a nest site poses additional demand, sometimes this can require more than one male and one female
- in these conditions it may be advantageous for fledglings to remain with parents, feed and defend the new brood until they gain skills to successfully establish a new nest
Helper-at-the-nest
larger the “clan” the better the nest site they can cooperatively defend
helpers’ short term postponement of their own reproduction is not a cost because it is unlikely that on their own the young offspring would survive
by assisting their parents during this time they are passing on their genes “by proxy”
Lions
female lions - social carnivory, they cooperatively hunt and cooperatively care for offspring so while many females can hunt many others can remain behind to care for cubs
male lions - compete to defend a pride of females, commit infanticide when females are not in estrus with the previous pride lion’s cubs still nursing
Reciprocal Altruism in Vampire Bats
vampire bats roost in large social groups
feeding is highly unreliable and high metabolisms make them susceptible to starvation and dehydration
at some point most individuals will require supplemental feeding from roost-mates
Conditions Favoring Reciprocal Altruism
low cost to giving aid, high benefit of giving aid
long-term association allows ample time to reciprocate and assess reciprocation
means of identifying individuals
means of punishing/exclusive non-reciprocators
Obligate Social Living
not just male and female - eg ants and bees
can lead to segregation of gender functions into more than 2 body types
1. drone (haploid male, fertile)
2. queen (diploid female, fertile due to added hormones)
3. worker (diploid female, sterile due to lack of hormones)
Honeypot Ant Eusociality
- workers act as storage vessels called “repletes” for honey used to feed brood and others
- specialized workers store honey in their abdomen for the colony to use when resources become scarce
- specialized body types are a form of “kin” altruism
Termite Eusociality
exception that proves the rule about kin selection
colony inbreeds generation after generation, resulting in very high levels of relatedness
have many non-reproductive “castes” with distinct body types and behavioral predispositions
gender roles are determined by hormones and pheromones as well as season, size, age of colony
fertilized females (queen gives birth to both males and females) will attempt to establish their own colony
Naked Mole Rat
a eusocial mammal
live in underground colonies, produced by a single breeding female who suppressed fertility of male and female workers with pheromones
a few breeding males also act as guards if colony is threatened
highly inbred, like termite colonies
workers dig elaborate tunnel systems and bring food back to the breeding chamber
Colugo “Flying Lemur”
closest "outgroup" to primates location: Southeast Asia not a primate or a bat, arboreal glider frugivory-insectivory uniparous, carries young
Tupaia “tree shrew”
next closest outgroup to primates arboreal locomotion insectivorous grasping hands, primate-like ears, binocular vision large snout/nose
Initial Primate Specialization 65 MYA
arboreal, noctural, insectivore/carnivore
large, forward-facing eyes
clinging-leaping-grasping
single birth, carries young
arboreal nocturnal hunting requires the ability to see in the dark and judge depth
Prosimians: Tarsiers
SE Asia, nocturnal hunter of insects, frogs, reptiles
clinging-leaping
solitary hunters, pair bonds and small groups
glaborous fingertips
Prosimians: Galagos (“Bushbabies”)
nocturnal, clingers and leapers
gumivory and insectivory
pair bonding with joint territory defense
multiparous litters
nest in hollow trees where young are left
Prosimians: Lemur Catta (“Ring Tailed Lemur”)
highly social, live in large multi-male multi-female groups, dominance hierarchies and male-male competition
tails are used for social communication
Lemur Diversity
typically diurnal, reduced eyes and enlarged snout with wet nose
include frugivores, folivores, and insectivores
include solitary and group living species
ancestors of lemurs were isolated on Madagascar with few predators or competitors and differentiated into diverse species with diverse adaptations
Prosimians
smaller bodies, smaller social groups, un-fused mandible, more insect and gum eating, nocturnal, some reliance on olfaction, limited color vision
Anthropoids
larger bodies, larger social groups, fused mandible, more fruit and leaf eating, diurnal, even less reliance on olfaction, extensive color vision
Locomotion
prosimians: clinging and leaping
monkeys and apes: arboreal quadrupeds, terrestrial quadrupeds, suspensory motion
Origin of the Anthropoid Primates
shift to diurnal feeding on fruit fruit co-evolved with birds as seed dispersers changes: 1. reduced eyes 2. change in tooth structure 3. reduced muzzle and nose 4. closure of the orbit
Loss of Vitamin C Synthesis
result of the shift to frugivory
- ascorbic acid not in diet = selection for endogenous synthesis
- dietary substitution
- dependence and re-adaptation
Color Vision
fruit initially evolved for seed dispersal by birds, which have good color vision
ripeness indicated by color
dependence on dietary ascorbic acid associated with selection for 3-color vision, changes in taste receptors, tolerance for alcohol (from over-ripe fruit), etc
New vs Old World Monkeys
NWM - platyrrhine (outward facing nostrils), 3 premolars, can have prehensile tails
OWM - catarrhine (downward facing nostrils), 2 premolars, never have prehensile tails
New World Monkeys: Assortment
marmosets - gumivores, defend territory
owl monkeys - convergent adaptations with prosimians, only monkey adapted to nocturnal foraging
spider monkeys - locomotion using prehensile tail
howler monkeys - defend territory using loud vocalization aided by an enlarged lower jaw and hyoid bone
uakari - Ally Boville
Old World Monkey Specializations
bilophodont molars, ischial callosities
diverged from apes with respect to locomotor adaptations (quadrupedal monkeys vs suspensory apes)
OWM further diverged into two groups with respect to dietary adaptations and specialized digestion (folivorous colobines vs omnivorous cercopithecines)
Colobines
specialized group of Old World Monkeys with adapted digestive system for folivorous diet
arboreal quadrupeds
OWM Arboreal Quadrupedalism
equal length arms and legs, rigid lumbar spine, four grasping hands, horizontal body
Cercopithecines
omnivorous
includes baboons, macaques, guenons
terrestrial quadrupeds
OWM Cercopithecines: Macaques
cheek pouches allow them to carry extra food without eating it
live in multi-male, multi-female troops
males usually emigrate from their troop to a new one while females stay with kin
males often form small coalitions
female dominance inherited from mother’s rank
Advantages of Monkey Sociality
defense of patchy resources predator defense grazing tracking social transmission of foraging kinship continuity (of one sex)
Disadvantages of Monkey Sociality
sexual competition
feeding competition
male paternity uncertainty
Infanticide in Langur Monkeys
Hanuman <3 langurs live in large single dominant male polygynous troops
when the dominant male is deposed, the new male systematically attacks and kills infants
this brings females back into estrus
females may feign sexual receptivity in order to protect their infants
OMW Cercopithecines: Baboons
widespread in sub-Saharan Africa and live in large multi-male, multi-female groups that tend to forage on the ground
mandrills: baboon species that live in forests, form single-male polygynous groups and have been known to kidnap females from other groups; most sexually dimorphic baboons
gelada baboons: forage on grasses in very large troops of smaller harems controlled by highly aggressive dominant males
Southeast Asian and African Apes
apes are far less numerous than monkeys, range is restricted to tropical forests of Africa and SE Asia
reflects primary adaptations for aboreal locomotion and foraging
range of apes used to be significantly greater 10-20 MYA
Ape Locomotor Adaptations
lack of tail, long curved fingers, highly mobile shoulder girdle and hip joint, shortened thumbs, four grasping limbs, vertical posture, mobile lumbar vertebrae, conical rib cage, wrist rotation, scapulae (shoulder blades) located on back
Asian Apes: Gibbons
small frugivorous brachiators
specializations for brachiation, coupled with small body size, predispose gibbons to walk bipedally when on the ground
gibbon pairs cooperatively defend a territory by singing duets twice a day
each sex drives off rivals of the same sex
Asian Apes: Siamangs
like gibbons, sing to defend territory (but only males sing) and also pair bond
specialized throat pouches for long distance vocalization
Asian Apes: Orangutans
Borneo and Sumatra
sexually dimorphic, males distinguished by fat pads
locomotion - quadra-manual, suspensory, mixed terrestrial forms of locomotion
also: infant ventral clinging and dorsal riding
isolated foraging
Orangutan Terrestrial Locomotion
variety of knuckle-, fist-, and wrist- walking strategies
African Great Ape Habitats
African apes except humans are confined to the equatorial forests (this environment is not good for fossilization)
Gorillas and chimpanzee habitats overlap, but bonobos are separated by the Congo River
African Apes: Gorillas
- folivorous, enabled by large digestive systems
- most of the day is taken up with eating and moving
- single large dominant “silverback” male with 3-8 females, infants, and some adolescent males
- no distinct territories and forage over large areas
- mature males develop silver hair on their backs and defend troop females, while young males attempt to abduct females and form new troops
Gorilla Group Structure
single dominant, sexually dimorphic "silverback" male mobile female-group foraging female-defense polygyny female transfer (abduction) male emigration to form a new group
Gorilla Locomotion
Terrestrial: mostly knuckle-walking but some bipedal walking is possible
Arboreal: possible but limited by size, more common in females
African Apes: Chimpanzee
Pan troglodytes - Central and West Africa
omnivorous, arboreal and terrestrial foraging
knuckle-walking
large multi-male, multi-female social groups
male cooperative territory defense
male hunting of monkeys
Chimpanzee Foraging Patterns and Emigration
individual females and young forage together in male-defended territories
female foraging territories overlap with each other
females leave their natal troops at puberty to join nearby male territories, while males remain with their natal troops
Chimpanzee Male Kin-Coalitions
reduced male-male aggressive competition, male hierarchy weakly limits sexual access
female fertility is advertised by perineal swelling and active solicitation during estrus
male kin-cooperation: extensive repertoire of social communication, involving vocalization/grooming/tactile contact
Chimpanzee Group Structure
multi-male, multi-female groups
modest sexual dimorphism
territory defense by male coalition
female transfer/ male continuity
Chimpanzee Tool Use
transmission of tool use by observation and crude imitation; no clear evidence of “training”
mastering a complex skill may take many years
also seen in capuchin monkeys
made possible by combination of continued social living, need to extract difficult foods, hands, large brains
Chimpanzee Hunting and Violence
males engage in regular cooperative hunting of colobus monkeys
kills are shared with females and young
male groups may attack and kill isolated males from neighboring troops
infanticide of infants brought into territory may also occur
Chimpanzee Infanticide
female “troop transfer” occurs when she reaches young adulthood, may be pregnant at this time
males and females will often harass new females in the troop and can kill an infant she is carrying
females have also been known to kill infants of other females within the same troop
African Apes: Bonobos
Pan paniscus - Central African forests
omnivorous, forage in large multi-sex multi-age social groups
minimal dominance hierarchy, weakly territorial, highly sexual
bonobos do not overlap with chimpanzees and therefore are not in competition for the same foods
Bonobo Locomotion
versatile locomotion:
agile arborealists, and on the ground both knuckle-walking and bipedalism are common
foraging on the ground may require use of arms for carrying food and offspring while erect; infants usually ride dorsally
Bonobo Social Structure
mobile multi-male, multi-female foraging groups
territorial but with minimal territory conflict and common troop exchange, mostly females
minimal dominance hierarchy not predictive of mating priority, females often dominant over males, males inherit rank from mothers
Bonobo Sexual Behavior
sexual swelling in females is extensive and long-lasting and accompanied by intense interest; as a result, same-sex stimulation among friends is common
interactions may involve many individuals of any sex or age
why so frequent? food is widely available, not season, no hunting
no competition with gorillas, etc., for ground-based foods
minimal male-male competition, social groups are large and all individuals forage together
Chimpanzee vs Bonobo Sex and Social Function
chimpanzees: highly competitive male access, minimal male-female bonding, minimal same-sex copulation
bonobos: no sexual competition, strong male-female friendships, considerable same-sex copulation with friends
