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Outline Adaptation and Evolution Natural Selection Proposed by Darwin Natural Selection Sociobiology Kin Selection Reciprocal Altruism Parental Investment Sexual Selection Based on 3 fundamental observable facts: •Variation •Inheritance •Overproduction Reproductive Strategies of Male and Female Primates Adaptation and Evolution – processes of biological of change over time Variation - Lots of variation within and between species •Adaptation = short term change within a species (ex moth wings change color) (aka microevolution) •Evolution = long term change that results in the appearance of new species (aka macroevolution) How do species change over time? Inheritance - Variation passed from parents to offspring By a process called Natural Selection Natural selection is the primary mechanism of biological change over time (adaptation and evolution) Principles of inheritance were not known when Darwin wrote Origin of Species 1 Overproduction – there are always more offspring born than can or do survive to adulthood Survival & Reproduction Survival alone is not the important thing in understanding evolution by natural selection Survival and then reproduction is the key to understanding adaptation, and evolutionary change over time Fitness - The ability to live and reproduce Overproduction means competition to survive to adulthood Some will survive to adulthood Some won’t Who survives to adulthood? Those who happen to have variation (morphology or behavior) that helps them survive Who doesn’t? Those who have variation (morphology or behavior) that doesn’t help them survive The environment naturally selects some variants over others High fitness = high reproductive success Low fitness = low reproductive success Those traits possessed by individuals with high survival and reproductive success, will be passed to future generations at higher rates. The environment naturally selects some variations (some individuals) over others to survive and reproduce The population, over time, becomes better and better adapted to local environments. 2 Adaptation/Evolution by Natural Selection The differential survival and reproductive success in each parental generation (NATURAL SELECTION) leads to A change in the frequency of traits from one generation to the next (ADAPTATION/EVOLUTION) The traits that • are well suited to the environment • give individuals an advantage in survival and reproductive success The traits that • are not well suited to the environment • put individuals at a disadvantage in survival and reproductive success Will appear in increased frequency in future generations Will appear in decreased frequency in future generations Peppered Moth – nicely illustrates the following 4 points about natural selection and change 1. Evolution (change) operates on the population – no individual moth ever changed color. Individuals do not “evolve”. 2. Natural selection operates on the individual. Individuals survive and reproduce or not. 3. Variation had to be there in the first place 4. No such thing as an absolutely better variant – depends on the environment Adaptation/Evolution by Natural Selection The differential survival and reproductive success in each parental generation (NATURAL SELECTION) leads to Peppered Moth – example of natural selection producing adaptation A change in the frequency of traits in the next generation (ADAPTATION/EVOLUTION) Sociobiology • Applying evolutionary principles, specifically natural selection, to behavior • Trying to understand how behavior might be naturally selected/adapted/evolved • Asking how behavioral variations give individuals an advantage in survival and reproduction 3 Sociobiology Developed in the 1960’s and 1970’s with the development of 3 main ideas: 1. Kin selection 2. Reciprocal Altruism 3. Parental Investment It all started with the question of altruism – how could altruism evolve What is altruism? Popular definition - helping behavior, selfless behavior, sharing behavior. Opposite of selfish behavior. Biological definition - behavior that potentially improves the survival & reproductive success of the recipient while potentially endangering the actor Behavior that benefits the recipient at a cost to the actor • • • • Altruism and Group Selection Originally, altruistic behavior was thought to evolve because it was “for the good of the group” Group Selection – read in chapter 4 of text What was the problem with Group Selection? Based on what we know about natural selection, behavior that increases the survival and reproductive success of the actor will be passed on at higher rate to next generation (will be favored by natural selection) Behavior that decreases the survival and reproductive success of the actor will not So how could altruism possibly get passed on? How could it evolve? Altruism - examples The answer to this question was the origin of sociobiology Predator alarm calls Helping another monkey in a fight Defending a group member from a predator Feeding/caring for a baby Altruism evolves not by group selection, but by kin selection Kin selection first principle of sociobiology Hamilton 1963 & 1964 4 Kin selection based on a recognition that: Does altruism ever occur between non-relatives? • Each individual shares genetic material with their relatives Yes • It is possible to have some of your genes represented in the next generation even if you never reproduce, if your relatives are reproductively successful Altruism between non-relatives cannot be explained by kin selection Important new concepts introduced with kin selection Reciprocal Altruism 1. Natural selection can operate on genes rather than individuals 2. Inclusive fitness – individual fitness plus effect upon the fitness of relatives Read about Hamilton’s model in Chapter 4 Food sharing Most altruism in primates does occur between kin Coalition formation supporting your kin in agonistic encounters Alternative explanation proposed: Reciprocal Altruism (Trivers 1971) Helping behavior between non relatives Individuals help others so that they will get help in the future when they need it. Eventually it will pay off – benefits to the individual will outweigh the costs Review Sociobiology Social grooming Developed in the 1960’s and 1970’s with the development of 3 main ideas: 1. Kin selection 2. Reciprocal Altruism 3. Parental Investment It all started with the question of altruism – how could altruism evolve 5 Parental Investment (Trivers 1972) Sexual Dimorphism Kin selection and reciprocal altruism are about explaining altruism Parental Investment is about explaining sex differences in behavior (& morphology) Parental investment is a reformulation of Darwin’s theory of Sexual Selection Sexual Dimorphism Primates Sexual Selection – a mechanism proposed by Darwin to explain secondary sexual characteristics What are secondary sexual characteristics? •features of sexual dimorphism •male female distinctions that appear at puberty •distinctions between adult males and females that go beyond basic reproductive differences Why was another mechanism needed to explain these features? Because secondary sexual characteristics are 1. Not easily explained by natural selection – – •differences in size, coloration, weaponry, shape, ornamentation not apparently beneficial to survival sometimes even detrimental to survival 2. Secondary sexual characteristics don’t appear until adulthood 6 According to Darwin, secondary sexual characteristics evolved due to Male male competition or Female choice Male – male competition (intrasexual selection) Secondary sexual characteristics give males a reproductive advantage through competition with other males. Natural Selection (NS) vs Sexual Selection (SS)? Reproductive success is the bottom line for both, but • NS emphasizes survival where SS cuts straight to reproduction • NS emphasizes getting to adulthood (the regular season). Once there, SS kicks in (the playoffs). • The “environment” is different – for SS it is often a social environment doing the selecting • SS operates more on males Triver’s (1972) Applied the principles of sexual selection to male-female behaviour as well as morphology Identified something he called Parental Investment. Trivers states that "the relative parental investment of the sexes in their young is the key variable controlling sexual selection" Female choice (intersexual selection) Secondary sexual characteristics give males a reproductive advantage by making them more attractive to females. Definition of Parental Investment: "Any investment by the parent in an individual offspring that increases the offspring's chance of survival (and hence reproductive success) at the cost of the parent's ability to invest in other offspring" 7 Any investment that a parent makes in one offspring decreases the ability to invest in others Limited “investment” budget – spending in one place means not spending in another The different initial and then subsequent levels of parental investment leads to very different strategies for males and females (main thrust of Triver’s famous 1972 paper) Females – be very choosy – choose males that will provide assistance, good territory, good genes etc. Males – go for as many fertilizations as possible and move on – little or nothing to lose Great potential for conflict and competition between Variation between and within species 1. parents and offspring (weaning conflict) 2. successive offspring (sibling rivalry for care/attention) The more similar the level of parental investment between the sexes, the more similar the sexes will be in behavior and/or morphology Examples: • Monogamy and monomorphism (owl monkey) • Polygyny and dimorphism (hamadryas baboons) • Within species – human mate choice studies 3. Parents (battle of the sexes) Important point – it is not about being male or female, per se, but about the level of PI – this is what determines sex differences Typical* Male vs Female PI in Mammals Females HIGH • Large nutrient rich egg • 1 of approx. 400 • Gestation • Lactation Males LOW • Tiny sperm (DNA with tail) • 1 of approx. 4 billion • no • no 3 reasons that male and female reproductive strategies and behavior are typically so different: 1. Different levels of parental investment 2. Different variance in reproductive success (aka lower potential reproductive output for females) 3. Different limiting factors * Lots of variation between and within species 8 Variance in reproductive success among female primates: Females typically have similar numbers of babies They typically have X number of offspring where X = reproductive lifespan divided by interbirth interval. 3. Limiting Factors Females limited by access to energetic resources Males limited by access to reproductive opportunities Not that much difference between least and most successful females (at least not compared with males) Variance in reproductive success among male primates: Much more variation among males than among females Males could be big winners or big losers. One male could theoretically be responsible for 100% of the offspring born. Or none. The stakes are much higher for males, and the reproductive striving therefore much more intense. Review – 3 factors responsible for different behavior/reproductive strategies of male and female primates: Females • Less variance/lower total output • More parental investment • Limited by access to energetic resources Males • Great variance/higher total output (potentially) • Less parental investment • Limited by access to fertile females Guinness World Book of Records 1997 Maximum number of offspring produced by a single human female = 69 Maximum number of offspring produced by a single human male = 888 Much greater variance in reproductive success for males than for females (at least in theory) Much greater potential reproductive output for males than for males 9