Day 15 october 28th chapter 9

Editor's Notes

• #4: The grasslands of the western United States are home to the Belding’s ground squirrels. They form large colonies of hundreds or even thousands of individuals. Because the colonies are so large, they attract many birds of prey. When a bird comes to the colony, it succeeds in killing a squirrel about 10% of the time. Squirrels have a system for reducing predation risk, however, that resembles a neighborhood watch program. When an aerial predator approaches, it is common for a squirrel standing on top of a burrow to produce a loud whistle-like call that serves as an alarm, warning other squirrels. Upon hearing an alarm call, squirrels quickly take cover in their burrows. Making an alarm call is a very dangerous activity: about half the time that an alarm call is made, the squirrel making the alarm call is attacked by the predator (Figure 9-9 Protecting relatives by making an alarm call). This raises the question: why would any squirrel make an alarm call? And given that some do, which squirrels are most likely to engage in this altruistic-appearing behavior? It certainly isn’t random: About 80% of the squirrels making alarm calls are female. Moreover, among females, older females are five times more likely than young females to make alarm calls.   These differences between male and females, and between young and old, reveal that alarm calling is about protecting relatives. The more kin an individual is likely to have, the more likely that individual is to call. Because males disperse long distances to new colonies shortly after reaching maturity, most adult males don’t live near their parents, siblings, or other relatives besides their own offspring. Females on the other hand, remain near the area where they were born and are likely to have many close relatives nearby. Older females, in particular, are likely to have the largest number of relatives.  
• #5: The biologist W.D. Hamilton expressed the idea of “kin selection”—that one individual assisting another could compensate for its own decrease in fitness if it helped a close relative in a way that increased the relative’s fitness. Altruistic-appearing behaviors will most likely occur when the benefits to close relatives are greater than the cost to the individual performing the behavior. The more closely related two individuals are, the more likely they are to act altruistically toward each other.  
• #6: Belding’s ground squirrels probably don’t know exactly how closely they are related to the squirrels around them, so it is likely that they use rules of thumb. In a clever experiment that tested this idea, researchers trapped an adult female and relocated her far away in a ground squirrel colony in which she had no close relatives. If she were to determine exactly how closely she was related to the squirrels around her, in this situation it would not behoove her to make an alarm call. What the researchers found, though, was that she was just as likely to sound the alarm as if she were on her home territory, surrounded by close relatives. It appears that females use a rule-of-thumb that says “If I am an older female, behave as if I have many close relatives around me.”   Figure 9-10 Following simple behavioral rules. A transplanted female Belding’s ground squirrel that is not related to the squirrel colony behaves as though she is.
• #7: One way of analyzing altruism is to observe how people give their money and property away when they die. Kin selection theory gives rise to the prediction that people will bequeath their assets based on their degree of relatedness to others. In an analysis of 1,000 randomly chosen wills, researchers tested two predictions of kin selection: 1) that individuals would leave more of their estates to genetic kin (and to spouses, who would distribute their assets to genetic kin), and 2) that closer relatives would be left greater amounts than more distantly related individuals.
• #8: Both predictions were supported: 92% of assets divvied up in the wills were given to genetic kin. 46% was given to siblings and children. Only 8% was given to half-siblings, grandchildren, and nieces and nephews. Less than 1% was given to cousins or more distant relations (Figure 9-11 Reading the will).
• #9: Based on the idea of kin selection, it is necessary to redefine an individual’s fitness. An individual’s fitness is not just his or her total reproductive output, which is called direct fitness. Fitness also includes the reproductive output that individuals bring about through their altruistic behaviors toward their close kin. This is called an individual’s indirect fitness. Taken together, the sum of an individual’s indirect and direct fitness is called inclusive fitness.  
• #10: No two individuals—with the exception of identical twins—are genetically identical. And because different individuals do not share all of the same alleles, we expect that they should experience some conflict. One disturbing example of such conflict takes place between a pregnant woman and her developing fetus.   Specifically, the mother and fetus differ when it comes to the question of how much food—doled out as nutrients in the blood flow across the placenta—the fetus ought to get. There is a point when it is in the mother’s best interests to reduce the amount of glucose and other nutrients given to the fetus. Her genes gain if she saves a bit for future fetuses, to whom she will be related equally.   Now consider the fetus’s “point of view.” Future siblings will carry some but not all of the fetus’s genes. Consequently, the fetus does not necessarily benefit from sacrificing nutritional intake for future siblings’ benefit. In a sense, it is a sibling rivalry that starts before the sibling is even conceived! The conflict results in a physiological battle throughout pregnancy. The fetus produces hormones that dilate the mother’s blood vessels, increasing the amount of sugar delivered to the fetus. In response, the mother produces insulin, which has exactly the opposite effect, reducing the amount of sugar in the bloodstream available to the fetus. In some mothers this conflict causes gestational diabetes—an inability to properly regulate her blood sugar levels—which disappears as soon as the baby is born. In all pregnancies, however, the conflict escalates until the mother is producing a thousand times the normal amount of insulin.
• #11: It is ironic that the study of altruism in the world reveals that natural selection has primarily produced selfish behavior among animals. And, as we saw in the previous section, when behavior appears altruistic, it frequently is because individuals are helping kin and, by doing so, advancing the copies of the genes they carry that happen to occur in their close relatives. In this section, we explore the conditions that give rise to reciprocal altruism, how it may have arisen, and why it is so common among humans while so rare among most other animal species.
• #12: We’ll start by examining one species with well-documented altruistic-appearing behavior: the vampire bat. Vampire bats live in social groups of 812 individuals, roosting primarily in caves and hollowed trees. They feed by landing on large mammals such as cattle, horses, and pigs, piercing the skin with their razor-sharp teeth, and drinking the blood that flows from the wound. A chemical in their saliva keeps the blood from clotting, allowing them to feed for a longer period of time. Because of their small body size and high metabolic rate, vampire bats must consume almost their entire body weight in blood each night. If they go for more than about 60 hours without finding a meal, they are likely to die from starvation. Here’s where the apparent altruism comes in. A bat that has not found food and is close to death will beg a bat that has eaten recently for food. In many cases, the bat that has just eaten will regurgitate some of the blood it consumed into the mouth of the hungry bat, saving it from starvation. This act obviously has very high benefits for the recipient of the blood and carries a cost to the vomiting bat (Figure 9-12 The gift of life).
• #13: Kin selection is responsible for some of the blood sharing (females often regurgitate blood for their own offspring), but in many cases, individuals give blood to unrelated individuals. How might this behavior have arisen? To answer the question, it is important to note three other features of vampire bats. First, vampire bats are able to recognize more than a hundred distinct individuals. Second, bats that receive blood donations from non-relatives reciprocate significantly more than average. And third, bats that are not familiar with each other (and do not have a history of helping each other) generally do not regurgitate for each other.  
• #14: One method proposed to explain the evolution of this apparent altruism is that the bats “giving” blood to other bats in need are repaid the favor when they are in need of blood. Therefore, the act only appears selfless when in actuality it is selfish. With such reciprocal altruism, both individuals give up something of relatively low value in exchange for getting something of great value at a later time when they need it most. In other words, they are storing goodwill in another individual, much the way a person might put money in a bank for a rainy day. In both cases, individuals are protected from some of the world’s uncertainties.
• #15: Reciprocal altruism has also been documented in vervet monkeys. Monkeys that would otherwise be searching for food or, perhaps, a mate spend time grooming other monkeys (mostly by picking parasites from their fur). In return for their grooming behavior, the monkeys are more likely to receive assistance in response to their solicitations for aid (Figure 9-13 You scratch my back, I’ll scratch yours).
• #16: Reciprocal altruism can evolve if certain conditions are met:   1. Repeated interactions among individuals, with opportunities to be both the donor and the recipient of altruistic-appearing acts 2. Benefits to the recipient that are significantly greater than the costs to the donor 3. The ability to recognize and punish cheaters, individuals that are recipients of altruistic-appearing acts but do not return the favor In the absence of these conditions, selfishness is expected to be the norm among unrelated individuals. In the presence of all three conditions, on the other hand, reciprocity masquerading as altruism is likely to occur, as it does among the vampire bats.
• #17: As rare as it is in other species, reciprocal altruism is very common among humans. Friendship, among other human relationships, is built on reciprocity and is almost universal. The opportunity for friendship may be enhanced as a consequence of our long lifespan and our ability to recognize thousands of faces and keep track of cheaters. These features are essential in people engaging in reciprocal altruism because an individual becomes very vulnerable when she acts in an altruistic manner toward an unrelated individual. The risk is that the altruism will not be repaid, in which case the cheater enjoys greater fitness than the altruist.  
• #18: While cooperators, those who repay altruistic-appearing behavior, have evolutionary advantages over loners, this advantage disappears if the cooperators are the givers all or most of the time, never, or rarely getting anything in return. Thus, it pays to avoid cheaters, individuals who accept altruistic behavior without repaying it. The importance of keeping track of cheaters and of identifying good potential reciprocity partners (i.e., other cooperators) may be why humans are so interested in social information and seemingly trivial gossip.
• #20: Recall that fitness not only is an individual’s reproductive success, relative to that of other individuals in the population, but also depends on the specific environment in which the organism lives. An adult Belding’s ground squirrel that makes alarm calls generally increases her inclusive fitness by doing so because all around her are individuals who share her genes. The world in which alarm calling evolved did not include biologists with pick-up trucks who trapped female squirrels, drove them long distances, and released them into colonies of unrelated individuals. In a world where long-distance transplantation was common, alarm calling would not have evolved. People often find it difficult to maintain their body weight, craving high-fat food even as we know it may increase our risk of heart disease and shorten our life. Our ancestor’s hunter-gatherer world was characterized by unpredictable food sources that could not be stored for long periods of time. In response to this environment, humans evolved to have great appetites and taste preferences for calorie-laden foods. Food consumption and the storage of fat helped to protect individuals during times of starvation. Today, most of us have easy access to unlimited amounts of food, including fat-laden fast foods. Because humans have not yet adapted to this environment of plentiful food, our instincts to consume large amounts are strong, making it difficult for most people to control their body weight.
• #21: Let’s return to the donations to refugees on another continent. For 2 million years, our ancestors lived as hunter-gatherers in small groups of a few hundred people, at most. Their success depended on joint efforts against predators and prey; being “nice” paid off when the prospect of hunting alone or sleeping outside the camp meant almost certain death. The loners died, so we are descended from those who could work well with others. It is only recently, in the blink of an eye in evolutionary time, that humans invented agriculture, industrialization, and the means of food production and distribution. As a result of these changes, population group size has increased dramatically; on a given day, you may see 10 or even 100 times as many people as a hunter-gatherer ancestor might have seen. It is with this brain that you approach the issue of refugees halfway across the world, or a homeless person somewhere in the United States. It is almost certain that you will not have repeated interactions with these people. Nor will you ever be in a position to be helped by them. But in the world humans evolved in, such altruistic behavior would most likely have been reciprocated at some future time, and thus your instincts guide you to and reward you for your kindness. It feels right. Figure 9-14 Charitable acts can give us pleasure. In the small hunter-gatherer groups in which humans evolved, altruistic behaviors would be reciprocated. Today, the pleasure we feel in response to such behaviors remains—even though the favor may never be returned.
• #22: Kin selection and reciprocal altruism can evolve in a population of animals, as we have seen; and examples of kin selection abound in nature. As a consequence, casual observers of nature frequently see individuals acting in ways that appear altruistic, even though these individuals are truly acting—from the perspective of evolutionary fitness—in their own selfish interests. This nearly universal selfishness raises the question of whether evolution ever leads to behaviors that are good for the species or population but detrimental to the individual exhibiting the behavior, a process called group selection.
• #23: It might seem that evolution would favor individuals that behave in a manner that benefits the group, even it if comes at a cost to the individual’s reproductive success. But this does not happen. Imagine that a new allele appears in a population that causes the individual carrying the allele to double its reproductive output, even though this might spell doom for the species as individuals over-use their resources. An individual carrying this “selfish” gene will pass it to more offspring than an individual carrying the alternative allele that codes for the production of fewer offspring. And those selfish offspring also will pass on the selfish gene at a higher rate than the alternative gene is passed on. This scenario may lead to extinction of the species, yet it still occurs. With its market share perpetually increasing, the selfish gene eventually will predominate. Conversely, a “selfless” gene arising in a population in which all individuals carry the selfish gene will never increase its market share relative to the selfish alternative (Figure 9-15 Can a “selfless” gene increase in frequency in a population?).   Just because an outcome is best for the group doesn’t mean that natural selection will produce it. In the end, natural selection tends to cause increases in the alleles that benefit the individual carrying them, even when this comes at the expense of the group. In some special situations, it is possible for natural selection to lead to group selection. But the stringent conditions necessary for this to occur are so rarely found in nature that we almost never see it.
• #24: Section 9.3 Opener “Look at me!” The red-crowned crane (Grus japonensis) extends its wings during a courtship display, as potential mates look on.
• #26: The very definition of “male” and “female” hinges on a physical difference between the sexes. In species with two distinct sexes, a female is defined as the sex that produces the larger gamete, while a male produces the smaller gamete (Figure 9-16 Many sperm, just one egg). At conception, the mother’s material and energetic contribution to the offspring, the energy she will expend in the growth, feeding, and care of offspring—her reproductive investment—exceeds the father’s. This is true for all animals, whether mammals, birds, insects, or sharks. (It is also true for plants.) Not only are female gametes larger, they tend to be relatively immobile and produced in smaller numbers. Male gametes, on the other hand, while smaller, are more plentiful and very motile.
• #27: This discrepancy in size and quantity of sperm produced by a male as compared to eggs produced by a female may seem trivial, but it sets the stage for evolutionary developments that magnify this initial energetic difference in the reproductive investment.
• #28: For starters, the difference in the number of gametes that males and females can produce means that males have the potential to produce many, many more offspring than females. Put another way, a male’s total reproductive output, the lifetime number of offspring he can produce, tends to increase as the number of females he is able to fertilize increases. A female, on the other hand, does not generally increase her reproductive success by mating with additional males beyond the first (Fig. 9-17).
• #29: Two physical differences that can exist between males and females are particularly important when it comes to reproduction. First, in species with internal fertilization, which includes most mammals, fertilization takes place in the female. The offspring also grow and develop within the female’s body. The amount of energy females invest in reproduction is therefore much greater than males’ investment. Females’ reproductive investment also limits their reproductive output; They can be pregnant only once at a time. A second important physical difference between females and males occurs in the mammals: Lactation takes place in females and not in males (with a very small number of exceptions). In these species, then, nurturing during both pregnancy and lactation can be accomplished only by the female. This difference in reproductive investment has led to the evolution of some very different reproductive behaviors between males and females.
• #30: Although the gamete size difference is consistent across all animals (i.e., the egg is always bigger and energetically more costly to make than the sperm), the physical differences between the sexes in the early nurturing (i.e., gestation and feeding) of offspring can vary considerably across animal species. In some cases, male and female investments become more nearly equal after fertilization. In birds, for example, before emergence of the chicks, much of the development of the fertilized egg is external: the female lays an egg, but either the male or the female can protect and incubate the developing embryo. Further, birds do not lactate. Once hatched, the chicks must be fed—a task that can be done by both parents.
• #31: External fertilization in fish and amphibian species further reduces the reproductive investment of the female—she does not spend any energy as the fertilized eggs begin developing into embryos.
• #32: We have seen, then, that in species in which fertilization occurs inside the female body, there is an unequal energetic investment in reproduction, and as a result the female expends more energy in the growth and care of offspring (much more so for mammals than for birds). Another profound consequence of internal fertilization is that a male cannot be 100% certain that any offspring a female produces are his progeny. Because it is possible for a female to mate with multiple males, any of whom could be the father, male mammals and birds will always have some degree of paternity uncertainty.
• #33: The differing patterns of investment cause males and females to be vulnerable to exploitation at different stages of the reproductive process.
• #34: A study at Florida State University in 1978 and 1982 set up two different encounters for both males and females. 1. A person of the opposite sex said, “Hi. I have been noticing you around campus. I find you very attractive. Would you go out with me tonight?” 50% of men said yes; 50% of women said yes. A person of the opposite sex said, “Would you have sex with me tonight?” 75% of men said yes; 0 women said yes. Humans, like nearly all mammals, are characterized by greater initial investment by females. Consistent with this, females are much more discriminating, whereas males are less hesitant to take a mating opportunity. Figure 9-19 part 1 The choosier sex. Males and females differ in their attitude toward mating opportunities. For the female more than the male, the choice of the wrong mate could have expensive consequences.
• #35: For females, at the point of mating, the cost of a poor choice can have significant consequences from an evolutionary perspective—pregnancy and lactation, with offspring from a low-quality male or a male who deserts her and does not provide any parental investment (Figure 9-19 part 2 The choosier sex). For a male, the consequences need not be great, little beyond the time and energy involved in mating.
• #36: The fact that physical differences in males and females lead to differences in reproductive investment gives rise to two predictions about sex-related behavior: 1. The sex that invests more will be more discriminating. 2. The sex that invests less will compete amongst themselves for access to the higher-investing sex.
• #37: A dramatic illustration of how a high reproductive investment leads to the evolution of choosiness in mating behavior comes from the insect world. When bush crickets mate, the male loses about a quarter of its body weight contributing a massive ejaculate, which the female then uses for energy (Figure 9-20 A costly decision). Not surprisingly, male crickets are very choosy when selecting a mate. Their contribution during sex would be the equivalent of nearly 50 pounds of semen in humans. Male bush crickets reject small females that would produce relatively few offspring.
• #38: For males, we’ll see that the point of greatest vulnerability comes when they provide parental care to offspring. Due to paternity uncertainty, there is some chance that the male may be investing in offspring that are not his progeny. This is an action that has significant evolutionary costs: rather than increasing his own inclusive fitness, he is increasing the fitness of another male. Females, conversely, are not at all vulnerable at the point of providing parental care to offspring because a female can be completely certain that the offspring she cares for are her own.
• #39: Female choosiness (and the male-male competition it leads to) tends to increase the likelihood that a female will select only those males that have plentiful resources or relatively high quality genes, either of which is beneficial to the female, causing her to produce more or better offspring—where “better” may mean increased disease resistance or physical traits that will be found attractive by future mates. Female choosiness is manifested by four general rules.
• #40: 1. Mate only after subjecting a male to courtship rituals. In many bird species, including the western grebe, females require that a male perform an elaborate and time-consuming courtship dance before she will mate with him. This courtship dance involves fancy dives into water, graceful hovering, various head movements, and flamboyant twists and turns. The courtship process can go on for several days. But if the male passes the time-consuming audition he can generally be counted on to stick around to see a brood through hatching and weaning (Figure 9-21 part 1 Four factors that influence a female’s choice of mate).   Among bower birds, the courting males build a small thatched structure, at which they make displays that can involve complex dance steps and dramatic poses, while courting the females. Female birds will walk around the structure and inspect it. Only if they deem it satisfactory will they mate with the male. Males that are rejected by multiple females will usually tear down their bower and rebuild it from scratch.  
• #41: 2. Mate only with a male who controls valuable resources. Territorial defense is a common means by which males compete for access to females. Among arctic ground squirrels, for example, females choose to reside on territories defended by males and mate with the male on the territory in which they reside. With greater quality and quantity of resources in his territory, a male is better able to attract females, whose reproductive success is partly a function of the resource richness of the territory.   Figure 9-21 part 3 Four factors that influence a female’s choice of mate
• #42: 3. Mate only with a male who contributes a large parental investment up front. Better than a believable pledge to commit resources to his [okay? I changed her to his.] offspring is an actual on-the-spot exchange in which a female gets a male to actually give her his parental investment up front, in the form of resources that will help her maximize her reproductive success. In the hanging fly, for example, a female will not mate with a male unless he brings her a big piece of food—usually a dead insect. If a male offers a sufficiently large meal—called a nuptial gift, she will mate with him (Figure 9-21 part 2 Four factors that influence a female’s choice of mate). The larger the food item is, the longer she will mate, and the more she eats, the larger the number of eggs that she will lay. After about 20 minutes of mating, though, when a male has transferred all of the sperm that he can, he becomes likely to break off the mating, taking whatever remains of the “gift,” which he may use to try to attract another mate. Nuptial feeding is common among birds and insects.  
• #43: 4. Mate only with a male that has a valuable physical attribute. Male-male competition for the chance to mate with females can also take a more literal form: actual physical contests. Across the animal kingdom, from dung beetles to hippopotamuses weighing more than 5,000 pounds, male-male contests determine the dominance rankings of males. Females then mate primarily with the highest ranking males.   In a similar process, females sometimes choose not the best-fighting or largest males, but instead base their choice on some physical attribute such as antler size in red deer, the bright red chest feathers of frigate birds, or the elaborate tail feathers of the male peacock. In each case, the physical feature serves as an indicator to females of the relative male quality, possibly because the feature is correlated with the male’s health (Figure 9-21 part 4 Four factors that influence a female’s choice of mate).
• #44: 1. Mate only after subjecting a male to courtship rituals. 2. Mate only with a male who controls valuable resources. 3. Mate only with a male who contributes a large parental investment up front. 4. Mate only with a male that has a valuable physical attribute. Although it is generally the case that males compete for access to females, this is not the case for all species. In species in which males make a significant parental contribution (such as in the species in which males provide a nuptial gift), the males must be somewhat selective, too. And the greater the male investment, the greater the male choosiness. Figure 9-21 Four factors that influence a female’s choice of mate
• #45: If a male simply abandons a female following mating and searches for other mating opportunities rather than making any investment in the potential offspring, he has no risk of investing in offspring that are not his. This is one way to minimize the potential costs associated with paternity uncertainty.   This strategy, however, is not necessarily the most effective way for a male to maximize his reproductive success.
• #46: If a male has a hundred or even a thousand matings but no offspring survive, that behavior is not evolutionarily successful. Consequently, in species for which offspring survival can be enhanced with greater parental investment, there is an incentive for males to provide some parental care…
• #47: In situations in which males provide parental care, it is common for the male to reduce his vulnerability through some form of mate guarding. Mate guarding is a consequence of (and attempt to reduce) paternity uncertainty. As long as the offspring emerge from the female’s body, she can be certain that they contain her genes. In contrast, a male inhabits a “danger zone” that lasts as long as the female is fertile. If she mates with anyone else during this time, the offspring she produces may not be his. If he is going to help raise the offspring, he benefits by minimizing his risk in the danger zone. It is almost exclusively during this time period that mate guarding occurs.
• #48: If a male wants to ensure that a female does not mate with another male, why bother to stop mating at all? In many species, males take this approach to reduce risk in the danger zone. Among house flies, even though the male has completed the transfer of sperm to the female in 10 minutes of copulation, he does not separate from her for a full hour. Moths go even further and continue to mate for a full 24 hours. And in the extreme case of this strategy, certain frog species continue individual bouts of mating for several months (Figure 9-22 Preventing paternity uncertainty). If humans mated for a similar percentage of our lives, a single round of intercourse would last almost 10 years.  
• #49: In a slightly subtler form of mate guarding that occurs in reptiles, insects, and many mammalian species, males block the passage of sperm into the female by producing a copulatory plug. Formed in the female reproductive tract from coagulated sperm and mucus, copulatory plugs can be very effective. Male garter snakes that encounter female snakes with a copulatory plug, for example, do not attempt to court or mate with her, treating her instead as if she is not available.  
• #50: A much more extreme form of mate guarding occurs in the black widow spider: the male breaks off his sexual organ inside the female, preventing her from ever mating again. Interestingly, when the act is completed, the female usually kills and eats the male (Figure 9-23 A reasonable trade-off?). In sealing his mate’s reproductive tract, the male assures himself of fathering the offspring and in consuming her mate’s nutrient-filled body, a female gets resources that help her to produce the offspring.