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  1. Behavioral ecology emerged after Nikolaas Tinbergen outlined four questions to address when studying animal behaviors (Figure \(\PageIndex{2}\)) that focused on two levels of answers: What are the ultimate (evolutionary) explanations of behavior, and what are the proximate (physiological or developmental) explanations of behavior?

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  2. Describe different ways that species learn. Behavior is the change in activity of an organism in response to a stimulus. Behavioral biology is the study of the biological and evolutionary bases for such changes. The idea that behaviors evolved as a result of the pressures of natural selection is not new.

    • Overview
    • Key points
    • Introduction
    • Simple learned behaviors
    • Habituation
    • Imprinting
    • Conditioned behaviors
    • Classical conditioning
    • Operant conditioning
    • Learning and cognition

    Habituation, imprinting, classical conditioning, operant conditioning, and cognitive learning.

    •Habituation is a simple learned behavior in which an animal gradually stops responding to a repeated stimulus.

    •Imprinting is a specialized form of learning that occurs during a brief period in young animals—e.g., ducks imprinting on their mother.

    •In classical conditioning, a new stimulus is associated with a pre-existing response through repeated pairing of new and previously known stimuli.

    •In operant conditioning, an animal learns to perform a behavior more or less frequently through a reward or punishment that follows the behavior.

    •Some animals, especially primates, are capable of more complex forms of learning, such as problem-solving and the construction of mental maps.

    •Habituation is a simple learned behavior in which an animal gradually stops responding to a repeated stimulus.

    •Imprinting is a specialized form of learning that occurs during a brief period in young animals—e.g., ducks imprinting on their mother.

    •In classical conditioning, a new stimulus is associated with a pre-existing response through repeated pairing of new and previously known stimuli.

    •In operant conditioning, an animal learns to perform a behavior more or less frequently through a reward or punishment that follows the behavior.

    If you own a dog—or have a friend who owns a dog—you probably know that dogs can be trained to do things like sit, beg, roll over, and play dead. These are examples of learned behaviors, and dogs can be capable of significant learning. By some estimates, a very clever dog has cognitive abilities on par with a two-and-a-half-year-old human!1‍ 

    In general, a learned behavior is one that an organism develops as a result of experience. Learned behaviors contrast with innate behaviors, which are genetically hardwired and can be performed without any prior experience or training. Of course, some behaviors have both learned and innate elements. For instance, zebra finches are genetically preprogrammed to learn a song, but the song they sing depends on what they hear from their fathers.

    Learned behaviors, even though they may have innate components or underpinnings, allow an individual organism to adapt to changes in the environment. Learned behaviors are modified by previous experiences; examples of simple learned behaviors include habituation and imprinting.

    Habituation is a simple form of learning in which an animal stops responding to a stimulus, or cue, after a period of repeated exposure. This is a form of non-associative learning, meaning that the stimulus is not linked with any punishment or reward.

    For example, prairie dogs typically sound an alarm call when threatened by a predator. At first, they will give this alarm call in response to hearing human steps, which indicate the presence of a large and potentially hungry animal.

    Imprinting is a simple and highly specific type of learning that occurs at a particular age or life stage during the development of certain animals, such as ducks and geese. When ducklings hatch, they imprint on the first adult animal they see, typically their mother. Once a duckling has imprinted on its mother, the sight of the mother acts as a cue to trigger a suite of survival-promoting behaviors, such as following the mother around and imitating her.

    How do we know this is not an innate behavior, in which the duckling is hardwired to follow around a female duck? That is, how do we know imprinting is a learning process conditioned by experience? If newborn ducks or geese see a human before they see their mother, they will imprint on the human and follow it around just as they would follow their real mother.

    Conditioned behaviors are the result of associative learning, which takes two forms: classical conditioning and operant conditioning.

    In classical conditioning, a response already associated with one stimulus is associated with a second stimulus to which it had no previous connection. The most famous example of classical conditioning comes from Ivan Pavlov’s experiments in which dogs were conditioned to drool—a response previously associated with food—upon hearing the sound of a bell.

    As Pavlov observed, and as you may have noticed too, dogs salivate, or drool, in response to the sight or smell of food. This is something dogs do innately, without any need for learning. In the language of classical conditioning, this existing stimulus-response pair can be broken into an unconditioned stimulus, the sight or smell of food, and an unconditioned response, drooling.

    In Pavlov's experiments, every time a dog was given food, another stimulus was provided alongside the unconditioned stimulus. Specifically, a bell was rung at the same time the dog received food. This ringing of the bell, paired with food, is an example of a conditioning stimulus—a new stimulus delivered in parallel with the unconditioned stimulus.

    Over time, the dogs learned to associate the ringing of the bell with food and to respond by drooling. Eventually, they would respond with drool when the bell was rung, even when the unconditioned stimulus, the food, was absent. This new, artificially formed stimulus-response pair consists of a conditioned stimulus, the bell ringing, and a conditioned response, drooling.

    Operant conditioning is a bit different than classical conditioning in that it does not rely on an existing stimulus-response pair. Instead, whenever an organism performs a behavior—or an intermediate step on the way to the complete behavior—it is given a reward or a punishment. At first, the organism may perform the behavior—e.g., pressing a lever—purely by chance. Through reinforcement, the organism is induced to perform the behavior more or less frequently.

    One prominent early investigator of operant conditioning was the psychologist B. F. Skinner, the inventor of the Skinner box, see image below. Skinner put rats in boxes containing a lever that would dispense food when pushed by the rat. The rat would initially push the lever a few times by accident, and would then begin to associate pushing the lever with getting the food. Over time, the rat would push the lever more and more frequently in order to obtain the food.

    Not all of Skinner's experiments involved pleasant treats. The bottom of the box consisted of a metal grid that could deliver an electric shock to rats as a punishment. When the rat got an electric shock each time it performed a certain behavior, it quickly learned to stop performing the behavior. As these examples show, both positive and negative reinforcement can be used to shape an organism's behavior in operant conditioning. Ouch! Poor rats!

    Operant conditioning is the basis of most animal training. For instance, you might give your dog a biscuit or a "Good dog!" every time it sits, rolls over, or refrains from barking. On the other hand, cows in a field surrounded by an electrified fence will quickly learn to avoid brushing up against the fence.4‍ 

    Humans, other primates, and some non-primate animals are capable of sophisticated learning that does not fit under the heading of classical or operant conditioning. Let's look at some examples of problem-solving and complex spatial learning in nonhuman animals.

  3. Behavior is the change in activity of an organism in response to a stimulus. Innate behaviors have a strong genetic component and are largely independent of environmental influences. In other words, these instinctive behaviors are “hard wired.”

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  5. One goal of behavioral biology is to dissect out the innate behaviors, which have a strong genetic component and are largely independent of environmental influences, from the learned behaviors, which result from environmental conditioning. Innate behavior, or instinct, is important because there is no risk of an incorrect behavior being learned.

  6. The biology of behaviour: scientific and ethical implications. The human brain is the most complex of all biological organs; it not only gives rise to consciousness—that most fascinating but elusive phenomenon—but also mediates our behavioural responses. The structure of the brain and its higher cognitive functions are the product of ...

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