Transcript Slide 1
Family groups – Conflicts and Interests among the Family - Before birth: Optimal sex ratios and adjusting for the circumstances - Conflicts between parents and offspring - Conflicts among siblings Males produce lots of sperm and a few males monopolize mating Male Female 50 Elephant seal Red deer Man Percent of copulations 100 24 888 8 14 69 0 1 2 3 4 5 6 7 8.............14 If its true, why not produce a sex ratio of say 20 females for every male? and do most organisms deviate from 1:1? However, the argument should really be stated in terms of parental investment and not reproductive success...b/c what if sons are 2x as costly to produce? Why might there be gender differences in the cost of offspring? FL scrub Jay In cooperative breeding birds, it is often males who stay behind to help and sex ratios are malebiased Local Mate Competition – the “Wasted Son” Suppose 2 sons compete to mate with the same female. An extreme case, but it illustrates that when brothers compete for the same females one male is wasted from the mother’s point of view An extreme example is a totally inbred population because all daughters are fertilized by the sons. What to do? Just enough sons to mate with all females - Acaronphenox mite – male are never born, rather they mate with sisters while still in the mother and dies. There broods consist of about 20 females and one male Trivers-Willard Effect An ESS 1:1 sex ratio is at the population level. Individuals may specialize (1:5 sex ratio) if an an equal number of other individuals specialize at 5:1 ratio In more or less monogamous human populations, there is evidence that women tend to marry up the socioeconomic scale. As a consequence, some women on the upper end of the scale would be left w/o men to marry while the same would be true for men at the opposite end of the scale Since there is a strong tendency for the socioeconomic status of the parents to determine the socioeconomic status of the children.... ... women at the upper end should produce sons while women at the lower end should produce daughters In animals.... (1) Population sex ratio must be 1:1 so that specialization in sons is cancelled by reciprocal specialization in daughters (2) One sex exhibits higher variance in Rep. Success than the other Male Female Elephant seal Red deer Man 100 24 888 8 14 69 (3) Parents can assess – consciously or not (“inherited”) – whether their offspring are likely to fall at the upper or lower bound of Rep. Success distribution e.g., in the above example, if you know your offspring are likely to succeed as competitive elephant seals, you will have more grandchildren if you have sons sons daughters daughters have higher success sons have higher success Sex ratio of offspring is biased toward males when females are high ranking Sex ratio of offspring is biased toward females when females are low ranking An experimental test: Virginia Opossum (Austad and Sunquist 1986) Experimental mothers were fed sardines during during the breeding season and gestation period, while controls received no food supplementation supplemented young had greater mass and male biased sex-ratio Possible mechanisms leading to biased sex ratios In Humans: 1,014 random males of the US èlite born between 1860-1939 - 1,180 sons and 1064 daughters - ratio 1.11:1 (differs from US mean of 1.06) 1,757 male of the German èlite - 1,473 sons and 1,294 daughters - 1.138:1 (differs from German mean of 1.05) 1,179 males of the British èlite - 1,789 sons to 1,522 daughters - 1.1754 (versus 1.06 mean) Ulrich Mueller (1993) Gaulin and Robbins (1991) Trivers-Willard Effect in contemporary NA Society Data derived from a National questionnaire, N=906 100 M Percent of older children breast fed 80 F 60 40 < 10K > 60K 1550 Interval (days) before younger child M 1450 1350 1250 F 1150 < 10K > 60K Duration of breast-feeding (mo.) Percent of older children breast fed 10 100 Interval (days) before younger child 2400 F 80 8 F M 60 2000 M 6 1600 M 4 40 Absent Present Male 1200 Absent Present F Absent Present How do they do it- Part II As Trivers has noted,the sex ratio at birth is only one indication of how parents distribute their investment (or alter their sex ratio) among male and female offspring Dickemann (1979) has argued that historical preferential female infanticide among high-status families of Europe and Asia reflects T-W investment bias Similarly, Voland (1988) has shown among 17-19th century German households a male based child mortality among all classes except the land-holding class Boone (1988): 15-16th century Portuguese nobility – highest classes invested more into sons (via estates), whereas the lower classes into daughters (via dowry) And on and on and on.... Parent-offspring conflict Classically viewed from perspective of parents Parents allocate investment to their young so as to maximize the number of surviving young, (or max LRS) and one imagines the offspring as passive vessels Once we see the offspring as an acting participant wanting to maximize its RS we realize that the offspring will presumably want more investment from the parents than the parents are selected to give Herein lies the conflict.... Fundamental Tradeoff between present and future reproductive effort, such that too much effort to the present compromises a parent’s survivorship leading to early death.... offspring survival rate parent parental investment What’s true about future RS if you die early? If the mother has fewer future offspring, is there a fitness cost to the present offspring? Tradeoffs among the offspring and among the parents... Parents – if I give too much PI now I will have fewer young in the future Offspring – if I demand too much PI now I will have fewer siblings in the future The key to seeing the conflict between parents and offspring is The offspring is related to itself by r = 1.0 and to its future siblings by r=0.50 (full sibling) or r = 0.25 (half sibling) Offspring should be selected to demand PI until the cost to the mother is twice the benefit to itself (or 4 times if half-sibs) because the mother’s future offspring are devalued by the current offspring (2) Weaning and begging In birds and mammals, young are “weaned” off a diet almost exclusively provided by the parents. There is a decline in food delivered to chicks as begging and contact-seeking behavior increases in many birds and mammals. Often, out-right aggression is use as the last act that dissolves the family. Time course of Parent-Offspring Conflict 2 B/C of PI No conflict here, adaptive to provide PI from either’s perspective No conflict here, adaptive to cease PI from either’s perspective 1 time conflict here; parents selected to stop offspring selected to be demanding Offspring as psychological manipulators Obviously young can not fling their mother to the ground to nurse at will... Presumably the young have better knowledge of their condition and need to communicate this to the parents. Both parties benefit from the communication, but almost immediately this system is subject to manipulation. (3) In-utero conflict in humans and mammals “inside the mother the offspring is expected to employ chemical tactics [to compete effectively with its parent]” (Trivers 1974) In-utero, there is potential conflict if the amount and duration of PI can be influenced by genes expressed in the offspring. In placental mammals, the fusion of fetal and maternal organs or tissue for the basis for physiological exchange Crespi and Semeniuk (2004) proposed that constrained antagonistic coevolution between parents and offspring in traits that influence PI has resulted in extreme diversity of placental mammals... There is no other mammalian organ whose structure and function are so species diverse as those of the placenta. This is curious since the ”purpose” of the placenta, presumably, is the same in all species (Faber et al. 1992) Uterine Tug-of-war: (1) Increased fetal trophoblast “invasion” with the uterine lining (2) Counter increased invasiveness via at least 3 mechanisms: - maternal secretions to reduce invasion, including stronger maternal immune response - evolution of stronger maternal epithelial barriers - shedding of overly invasive trophoblasts with uterine lining Interestingly, transplanted trophoblasts (in mice and pigs) to nonreceptive regions of the uterus results in enhanced trophoblast invasion, which suggests this process is usually suppressed in normal pregnancy (3) Fetal tissue (in humans) and estrogens (horses) function to increase utero-placental blood flow (via dilation of vessels), thereby supplying the fetus with more resources. Suppression of these estrogens in horses leads to smaller foals. Pre-eclampsia and gestational diabetes