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Hank: Behavior is action in response to a stimulus. My cat Cameo is now responding to both an external stimulus, the sound of a bag of treats, and an internal stimulus, her hunger, or at least her insatiable desire for treats. (lively music) Sometimes animal behavior can seem kind of far out. But if you look closely enough you can see how all behavior serves a purpose to help an animal mate, eat, avoid predators, and raise young. And since behaviors can come with advantages like these, natural selection acts on them, just as it acts on physical traits ensuring the success of animals who engage in beneficial behaviors while weeding out those that do stupid, dangerous, or otherwise unhelpful stuff. The most beneficial behaviors are those that make an animal better at doing the only two things in the world that matter. Eating and sexing. Still, that doesn't mean that all behavior is just about looking out for number one. It turns out that some advantageous behavior is pretty selfless, actually. More on that in a minute. But first behavior is really just a product of a pair of factors. Morphology or the physical structure of an animal and physiology or the function of that morphology. Now an animal's behavior is obviously limited by what its body is capable of doing. For example, Cameo does not have opposable thumbs, so as much as she would like to get into the treat bag by herself, she cannot. This limitation is strictly hereditary. No cats can open treat bags with their thumbs because no cats have opposable thumbs. Though some cats do have thumbs. It's the same way that a penguin can't fly to escape a predator, or a gazelle can't reach the same leaves as a giraffe can. Similarly, behavior is constrained by an animal's physiology. Like Cameo is built for chasing down little critters and eating meat, not beds of lettuce. This is because of her physiology, everything from her teeth to her digestive system are geared for eating meat and if she pounced on and ate every blade of grass she came across, let's just say I would not want to be in charge of that litter box. The traits that make up an animal's morphology and physiology are often heritable so we generally talk about selection acting on those traits. This natural selection hones these traits. It's really selecting their associated behaviors. It's the use of the trait, using wings and feathers to escape predators, or using a long neck to reach leaves that provides the evolutionary advantage. Still that doesn't mean that all behavior is coded in an animal's genes. Some behaviors are learned and even for animals that learn how to do things, natural selection has favored brain structures that are capable of learning. So one way or another most behaviors have some genetic underpinning and we call those behaviors adaptive. The problem is it's not always obvious what the evolutionary advantages are for some of the nutty things that animals do. Like why the snapping turtle always stick out its tongue? How does a tiny Siberian hamster find its mate miles across the unforgiving tundra? Why does a bowerbird collect piles of garbage? To answer questions like those we have to figure out one, what stimulus causes these behaviors, and two, what functions the behaviors serve. To do this I'm going to need the help of one of the first animal behavior scientists ever or ethologists, Niko Tinbergen. Tinbergen developed a set of four questions aimed at understanding animal behavior. The questions focus on how a behavior occurs and why natural selection has favored this particular behavior. Determining how a behavior occurs actually involves two questions. One, what stimulus causes it? And two, what does the animals body do in response to that stimulus? These are the causes that are closest to the specific behavior that we're looking at. So they're called the proximate causes. In the case of the male Siberian hamster the stimulus is a delicious smelling pheromone that the sexy female hamster releases when she's ready to mate. The male hamster's response, of course, is to scuttle surprisingly quickly over several miles, if necessary, to find and mate with her. So, the proximate cause of this behavior was that the girl hamster signaled that she was ready to knock boots and the male ran like crazy to get to the boot knocking. Asking the more complex question of why natural selection has favored this behavior requires asking two more questions. One, what about this behavior helps the animal survive and/or reproduce? And two, what is the evolutionary history of this behavior? These as you can tell are bigger picture questions and they show us the ultimate causes of the behavior. The answer to the first question, of course, is that the ability of a male hamster to detect and respond to the pheromones of an ovulating female is directly linked to his reproductive success. As for the the second question you can also see that male hamsters with superior pheromone detectors will be able to find females more successfully than other male hamsters and thereby produce more offspring. So natural selection has honed this particular physical ability and behavior over generations of hamsters. So who would have thought to ask these questions in the first place? And where is my chair? (piano music) Niko Tinbergen was one-third of a trifecta of revolutionary ethologists in the 20th century. Along with Austrains Karl Von Frisch and Konrad Lorenz, he provided a foundation for studying animal behavior and applied these ideas to the study of specific behaviors and for that all three shared the Nobel Prize in 1973. You may have seen the famous photos of young greylag geese following obediently in line behind a man. That was Lorenz and his experiments first conducted in the 1930's introduced the world to imprinting. The formation of social bonds in infant animals and the behavior that includes both learned and innate components. When he observed newly hatched ducklings and geese, he discovered that waterfowl in particular had no innate recognition of their mothers. In the case of greylag geese he found the imprinting stimulus to be any nearby object moving away from the young. So when incubator hatched gosslings spent their first hours with Lorenz, not only did they follow him, but they showed no recognition of their real mother or other adult in their species. Unfortunately, Lorenz was also a member of the Nazi party from 1938 to 1943 and in response to some of his studies on degenerative features that arose in hybrid geese, Lorenz warned that it took only a small amount of tainted blood to have an influence on a pure blooded race. Unsurprisingly, Nazi party leaders were quick to draw some insane conclusions from Lorenz's behavioral studies and the cause of what they called race hygiene. Lorenz never denied his Nazi affiliation, but spent years trying to distance himself from the party and apologizing for getting caught up in that evil. Now how exactly does natural selection act on behavior out there in the world? That's where we turn to those two types of behavior that are the only things in the world that matter, eating and sex having. Behavior associated with finding and eating food is known as foraging, which you've heard of. And natural selection can act on behaviors that allow animals to exploit food sources while using the least amount of energy possible. This sweet spot is known as the optimal foraging model. The alligator snapping turtle has optimal foraging all figured out. Rather than running around hunting down its prey, it simply sits in the water and food comes to him. So the alligator snapping turtle has a long pink tongue, divided into two segments, making it look like a tasty worm to a passing fish. In response to the stimulus of a passing fish it sticks out its tongue and wiggles it. Natural selection has, over many generations, acted not only on turtles with pinker and more wiggly tongues to catch more fish, it's also acted on those that best know how and when to wiggle those tongues to get the most food. So it's selecting both the physical trait and the behavior that best exploits it. And what could be sexier than a turtle's wiggly tongue dance? Well, how about sex? As we saw with our friend the horny Siberian hamster, some behaviors and their associated physical features are adapted to allow an animal to reproduce more, simply by being better at finding mates. Many times animals of the same species live close together or in groups and determining who in what group gets to mate creates some interesting behaviors and features. This is what sexual selection is all about. Often males of the species will find and defend a desirable habitat to raise young in and females will choose a male based on their territory. But what about those species, and there are many of them, where the female picks a male not because of that, but because of how he dances, or even weirder, how much junk he's collected? Take the male bowerbird. He builds an elaborate hut or bower out of twigs and bits of grass and then spends an enormous amount of time collecting stuff. Sometimes piles of berries and sometimes piles of pretty, blue, plastic crap. Ethologists believe that he is collecting the stuff to attract the female to check out his elaborate house. Once the female has been enticed to take a closer look, the male starts to sing songs and dance around, often mimicking other species, inside of his little house for her. Females will inspect a number of these bowers before choosing who to mate with. Now doing more complex dances and having more blue objects in your bower scores bigger with females. And ethologists have shown that a higher level of problem solving or intelligence in males correlates to both of these activities. So, yeah it took some brawn to build that bower and collect all that junk, but chicks also dig nerds who can learn dances, so the bowerbird's brain is evolving in response to sexual selection by females. This intelligence likely also translates into other helpful behaviors, like avoiding predators. So thanks to the evolution of behavior, we're really good at taking care of our nutritional and sexual needs, but what's confused scientists for a long time is why animals often look after others' needs. For instance vampire bats in South America will literally regurgitate blood into the mouths of members of its clan who didn't get a meal that night. How do you explain animals who act altruistically like that? We actually did a whole side show episode on this very subject, but basically we can thank British scientist William Hamilton for coming up with an equation to explain how natural selection can simultaneously make animals fit and allow for the evolution of altruism. Hamilton found that the evolution of altruism was best understood at the level of larger communities, especially extended animal families. Basically altruism can evolve if the benefit of a behavior is greater than its cost on an individual, because it helped the individual's relatives enough to make it worth it. Hamilton called this inclusive fitness, expanding Darwin's definition of fitness, which is basically how many babies somebody is making, to include the offspring of relatives. Now I guess the only question left is if I forget to feed you two, who's going to regurgitate blood into the other one's mouth? Yeah, there's probably a reason why that only happens with bats. (cat meows)