10.1: Learning, Genes, and Adaptation

Learning Objectives

1. Describe several definitions of learning which vary depending on theoretical perspective.
2. Discuss the claim that behavior is not random, but orderly and lawful.
3. Describe the fundamental law that governs the organization of behavior and the origin of that natural law.
4. Explain the similarities and differences between information in genes and learned information.
5. Discuss the nature-nurture continuum.
6. Describe examples of "biological preparedness" or biological constraints on learning.

Overview

In this section, we examine learning and how it contributes to the adaptive organization of mind and behavior. To understand learning within its broader biological context, it is important to compare it with genetic information, the other major source of information that organizes behavior into adaptive patterns. Learning refers to behavioral change as a result of experience, but learning is not a single, unitary phenomenon.  Instead there are many types of learning which are specialized to adapt to various features of the environment and to solve a variety of adaptive problems. In other words, learning evolved in many forms to serve a range of adaptive functions. Learned and genetic sources of information interact to organize behavior into adaptive patterns. The relative contribution of each varies with the species and the behavior in question. 

Learned and Genetic Sources of Information Shape Adaptive Behavior

Learning is defined differently by different psychologists depending upon the theoretical emphasis of the psychologist. Strict behaviorists rejected explanations of behavior that involved reference to internal mental processes. Instead, they emphasized the measurable relations between observable behaviors and observable stimuli. Consequently, behaviorists usually define learning as a change in behavior as a result of experience (excluding changes due to fatigue or other special circumstances). Cognitive psychologists, partially in reaction against strict behaviorism, are more likely to define learning as acquisition of information as a result of experience. From an evolutionary perspective, learning can be thought of as the acquisition of information during the lifetime of the individual animal. This last phrase differentiates learned information from genetic information which is acquired over the evolutionary history of the species.

Behavior is not random. Instead, like all of nature, behavior follows natural laws. An evolutionary perspective applied to learning emphasizes the role of learning as a means of achieving successful adaptation. Certainly, the ability to learn new behaviors in response to new situations encountered in the environment is valuable to survival and therefore increases the chances for reproduction and the transmission of one's genes into future generations, including genes for learning itself. This consideration is important because it suggests how and why learning evolved. It also suggest why behavior is orderly and lawful. Behavior is organized to promote survival and reproduction. This outcome is an inevitable product of the forces of evolution, most notably, natural selection. Thus, learning itself is an evolved property of nervous systems. The properties and laws of learning have evolved and are consistent with the primary law of evolution, natural selection. Learning reflects this law so that learned information organizes behavior in ways that serve an animal's biological fitness in the "struggle for existence" (to the extent that this is not true for any individual animal or human, we recognize some form of behavioral or psychological pathology). In section 10.2, we discuss in more detail how different forms of learning contribute to adaptation and improve biological fitness to particular features of the environment.

Figure \(\PageIndex{1}\): Dusky Langur infant and mother. In the first few months the infant is constantly breast-fed. The mother must teach her infant to eat leaves, as shown in this photo. (Image and caption adapted from Wikimedia Commons; File:Dusky Langur infant learning to eat leaves.jpg; https://commons.wikimedia.org/wiki/F...eat_leaves.jpg; by: Roughdiamond21; licensed under the Creative Commons Attribution-Share Alike 4.0 International license).

Behavior, and other traits of organisms, ultimately are organized by information. Just like a builder needs information in the form of structural plans to construct a building, information is required to build and operate an organism, including the guidance of its movement, its behavior. In short, behavior is not random, but orderly and lawful, because it is organized by information selected and filtered by two processes. Learning is one source of the information that organizes behavior into adaptive patterns. The laws of learning tend to organize learned behavior in ways that serve adaptation to the environment. This is because the laws of learning themselves are evolved properties of brains, and natural selection shaped these laws into adaptive form so that learned behavior tends to increase survival and reproduction. The second informational source for the adaptive organization of living things is in genes, already discussed briefly above, and in some detail in Chapter 3 on evolution and genetics. From earlier discussion, we know that genetic information is filtered over countless generations by natural selection and other processes of evolution (see Chapter 3) so that the information transmitted across generations in genes generates traits that, in general, contribute to survival and reproduction (genetic abnormalities are, of course, an exception). In short, information is required to organize living things and their functioning. This information comes from two sources: 1) genes and genetic evolution; and 2) learning and laws of learning (in those organisms that are capable of learning).

Let's compare these two sources of information which organize behavior into forms which help an animal adapt to environmental demands (see Table 10.1.1 below).

First, genetic information is available to all living organisms on earth, but learned information is available only to species whose brains are equipped with properties that make learning possible. Do flies learn? Do cockroaches? Do frogs? If they do, how much do they learn, how readily do they learn? How important is learning in the life of a frog compared to how important learning is in the life of a dog, a chimpanzee, or a human? In general, learning is more characteristic of the brains of the more complex animals, especially the mammals. Nevertheless, learning does occur even in many invertebrates (animals without backbones, such as insects or squid and octopus; I once visited the lab of a psychologist at the University of Hawaii who was studying learning in bees). Usually, however, the more complex the animal's brain, the more likely it is that the animal has well developed capacities for learning--capacities for the acquisition of information during its individual lifetime.

Secondly, genetic information has been acquired and transmitted generation after generation over the entire genetic history of the species. For example, think of flies, or pelicans. As you know from your own personal observation, flies are very good at flying. And so are pelicans. They ride the air currents perfectly and skim the ocean only inches above it, their paths rising and falling with the tops of breaking waves. As the wave rises they rise just enough to stay inches above the wave while flying at perhaps 30 miles per hour or more. When the pelican dives to catch a fish, it tucks its wings just at the right moment, and it rarely misses its target. How do these birds (and flies) know how to fly at all, and to fly and dive so well?

Figure \(\PageIndex{2}\): Brown pelican tucking its wings as it dives for a fish. Genetic information controls the complex, precisely timed movements involved (Image from Wikimedia Commons; File:Brown pelican (Pelecanus occidentalis occidentalis) diving.jpg; https://commons.wikimedia.org/wiki/F...is)_diving.jpg; By Charles J. Sharp, Sharp Photography; licensed under the Creative Commons Attribution-Share Alike 4.0 International license).

Over the evolutionary history of these diverse species (one an insect, the other a large bird), natural selection has preserved the genetic information that allows flies and pelicans to fly (they don't learn it); and it is just the right information needed to organize the complex movements of flight in these two very different species.

How did the "right" information, information that so perfectly organizes the movements required for flight, get into fly DNA and pelican DNA? Of course, the answer is natural selection. The wrong information has been weeded out over thousands of generations by natural selection, leaving for reproduction to future generations only the "correct" information, the information that guides with precision the complex movements required for flight in these two species. Note that this involves an information acquisition process, but it is species-wide and takes place over the thousands and thousands of generations that make up the genetic and evolutionary history of each species. The primary mechanism for acquisition of genetic information, the "correct" (i.e., adaptive) genetic information, is natural selection. Its method of transmission is heredity.

By contrast, learned information is acquired not over the genetic history of the species, but during the short lifetime of the individual animal. This is useful because some changes in the environment that may impact chances for survival and reproduction (successful adaptation) may be temporary or highly specific or even unique to the experience of only one or a few individuals within a species. Such changes are far too rapid and too specific to one or a few individuals for the relatively slow processes of natural selection to operate. For example, learning where the local water holes are would certainly be very useful to a lion during the dry season. Dependence upon genetics for this type of information isn't likely to work very well.  Learning mechanisms are required.

Figure \(\PageIndex{3}\): Lioness drinking from small water hole on the African savannah. Information about location of water on the savannah changes too rapidly to be incorporated into genes and therefore must be learned (Image from Wikimedia Commons; File:Lioness drinking.jpg; https://commons.wikimedia.org/wiki/F...s_drinking.jpg; By James Wagner; licensed under the Creative Commons Attribution-Share Alike 4.0 International license).

In sum, the mechanisms of information acquisition are different for genetic information (the law of natural selection and the laws of heredity) and for learning (the laws of learning; see below), but both result in information acquisition. In the healthy organism this information facilitates adaptation to the environment (with some exceptions as when learning goes awry; an example would be behavioral pathologies such as drug addiction where drugs such as amphetamine, by virtue of their chemical structure, activate pleasure circuits in the brain when they shouldn't be activated, thereby reinforcing harmful drug-taking behaviors; see chapter on psychoactive drugs).

Thirdly, the mode of coding and storage is different for genetic information compared to learned information. Remember, DNA forms a molecular code for coding and storage of genetic information. Learned information is coded and stored in memory systems in brains. We don't yet know all the details of the coding and storage mechanisms in learning. Early theories proposed a molecular storage mechanism in brain proteins. This led to memory transfer experiments in the 1970's wherein an animal was taught something, like a conditioned response to a signal, and then killed, and its brain ground up in a malt mixer; the liquid brain was centrifuged to extract brain proteins or RNA molecules, which were then injected into the brains of untrained recipient animals. These animals were then tested to see if they showed the conditioned response to the signal, that is, to see if they had memory for something they had not experienced themselves. Some positive results were reported by some labs; but most labs could not get the memory transfer effect and this line of research disappeared after a few years. Current research suggests that changes in existing synapses, making them more or less responsive, and/or formation of new synapses, stores learned information in memory.  We will cover the details of synaptic change in section 10.4.

Fourth, genetic information is transmitted across generations by genetic transmission (heredity). By contrast, learned information, in a few species including us, can also be transmitted across generations--not by genetic transmission, but instead by what is known as cultural transmission (tradition, imitation of the old by the younger members of the social group, books, teaching and learning, storytelling, and so forth). Cultural transmission occurs in relatively few species. We can do it; chimpanzees can do it to a limited degree; Japanese macaque (ma-cack) monkeys can do it to a limited degree. For example, one clever macaque female invented an efficient way to clean sand from wheat grains; other macaques in the group watched and learned, and the learned behavior spread throughout the group and then across generations--cultural transmission (Schofield, et al., 2018). However, rats, and most other species, can't do cultural transmission, and certainly no other species comes close to using cultural transmission to the extent that humans do.

Figure \(\PageIndex{4}\): Macaque monkey feeding. Learned behaviors can be transmitted across generations by cultural transmission by humans and a limited number of other species including macaques (Image from Wikimedia Commons; File:Artis Dining Japanese macaque (6807832054).jpg; https://commons.wikimedia.org/wiki/F...807832054).jpg; By Kitty Terwolbeck; licensed under the Creative Commons Attribution 2.0 Generic license).

Cultural transmission takes a special kind of brain but most species don't have the kind of brain circuit organization required for cultural transmission (the transmission of learned information from generation to generation). Language in humans, of course, plays a crucial role in human cultural transmission, but so do technological and social inventions such as writing, the printing press, film, video, formal education, research institutions, computers, and the internet. All have magnified human cultural transmission and played an enormous role in recent human adaptation (Koenigshofer, 2011).  Just as swimming can be considered a defining adaptation of fish, cultural transmission is an adaptive specialty of humans.

 

                                                 Period of Acquisition                Laws of Acquisition         Mechanisms of Encoding/Storage    Transmission Mechanism

Genetic information	acquired over generations (slow information acquisition process), producing species-wide, innate adaptations	acquired by the laws of evolution, primarily by natural selection, over the evolutionary history of the species	encoded and stored in a molecular code in DNA in the genes and chromosomes of the nucleus of cells	transmitted across generations by genetic transmission (heredity, inheritance)
Learned information	acquired during an individual's lifetime (fast information acquisition process) producing rapid and individualized adaptation	acquired by the laws of learning such as the law of effect and association by occurrence of events close in time	encoded and stored in changes in neural circuits probably involving changes in synaptic connections between neurons	transmitted across generations by cultural transmission

Table 10.1.1.  Comparisons between genetic and learned information organizing behavior into adaptive patterns.  Adaptations, including behavioral adaptations, are not random, but highly structured. This structure requires information. Two sources of information organize behavioral adaptations--genetics and learning, nature and nurture, in interaction (table and caption by Kenneth A. Koenigshofer, PhD; licensed under the Creative Commons Attribution-Share Alike 4.0 International license).

Types of Biological Adaptation

At this point it is useful to note that biological adaptations can be loosely categorized into three (overlapping) general types:

1. Anatomical adaptations (structural features of the organism such as having fur, or wings, or fins, or hands, or bones, or a liver, or a large brain with a well developed cerebral cortex). These are organized by genetic information and change (evolve) only relatively slowly (although small changes, for example, in beak size and thickness of finches in the Galapagos, can occur much more rapidly; Grant & Grant, 1993).
2. Physiological adaptations (internal dynamic processes of the organism such as photosynthesis in plants, digestive processes in animals, the immune system's operations, shivering in response to cold, sweating in response to excess heat, the circulatory system's operations, etc.). These are organized by genetic information and change (evolve) only relatively slowly, although the immune system shows significant plasticity during the individual lifetime to deal with newly encountered pathogens.
3. Behavioral adaptations(the goal-directed movements of the organism and the mental processes such as thoughts, plans, emotions, perceptions, reasoning processes, imagination, etc. that underlie and control these movements). As implied above, behavioral adaptations can be organized by genetic information (genetically preprogrammed, "instinct" and reflexes; flying in flies; swimming in fish; feeding in frogs; sex drive in humans) or by learned information (acquired during the lifetime of the animal), or by a combination of both of these sources. It is important to be aware that the abilities for learning in any species are themselves genetically evolved. Species that can learn can do so only because learning, and the properties of the nervous system that make learning possible, evolved in those species, including humans, as a result of natural selection.

Figure \(\PageIndex{5}\): Human sex drive is inborn. It can be thought of as a psychological adaptation involving intense emotions leading to reproductive behavior within the context of a pair-bond, increasing likelihood of surviving offspring. Learned cultural practices influence courtship practices and the expression of innate sex drive. (Image from Wikipedia Commons; File:Family love wiki008.jpg; https://commons.wikimedia.org/wiki/F...ve_wiki008.jpg; By Shagil Kannur; licensed under the Creative Commons Attribution-Share Alike 4.0 International license).

Behavioral adaptations can be categorized, at least conceptually, into 2 major types based on the source of the information that controls them: 1) learned adaptive patterns of behavior, often called "adaptive adjustments" by biologists (organized primarily by learned information, that is, information acquired during the lifetime of the individual animal, such as agricultural practices in humans) and 2) genetic or innate behavioral adaptations (organized by hereditary information, information stored in DNA, acquired over the evolutionary history of the species). However, many behavioral adaptations (perhaps most, in the more complex animal species especially) are a combination of both sources of information. Learning always involves genetic information to some degree because even "general" forms of learning such as conditioning incorporate and interact with information acquired by genetic evolution.  Ability to learn is genetically evolved and learning processes follow laws of learning evolved by natural selection.  Even conditioning mechanisms incorporate genetically encoded information that guides learning (Chiappe & McDonald, 2005; Gallistel, 1992, 2000; Koenigshofer, 2017).

Some authors refer to learned behaviors as "adjustments," rather than "adaptations." However, here we sometimes may use "learned behavioral adaptations" to emphasize that learning capacities themselves are genetically evolved, and that, generally (in healthy animals), learning and learned behaviors typically contribute to improved adaptation because of the evolved adaptive organization of pleasure and pain circuitry in the brain. For example, reward mechanisms in the brain that reinforce learned voluntary behaviors involve genetically evolved reward circuits, and punishments that inhibit voluntary behaviors involve activation of genetically evolved pain and fear circuits (reduced activity in reward circuits can also be "punishing"). In addition, behavioral change in general is a consequence of other evolved properties of brains at the synaptic level (see Section 10.4). The behavioral guidance provided by these circuits improves adaptation, increasing biological fitness (chances of survival and reproduction).

The behaviors of various animal species can be thought of as falling on a nature-nurture continuum, with some behaviors in some species (flies, roaches) being almost completely at the nature (innate) end of the continuum (i.e. behavior determined by genes and genetic evolution), while at the other end, the nurture end of the continuum, are behaviors which are dependent primarily on information acquired during the lifetime of the individual (i.e. learned information), such as termite fishing by a chimpanzee, or in the case of humans, how to make a flint spearhead, how to make a light bulb, do calculus, dig a well, or engage in modern agricultural practices (Koenigshofer, 2011, 2016). See figure 10.1.6 below.

Long-Term, Across-Generation Categorical Information vs. Short-Term Specific Information

One way to look at learning and genetics involves a general principle governing these two sources of information. Recall from Module 3.1 of this text that natural selection can only act on long-term, recurrent, across-generation environmental conditions in order to create complex adaptations. To illustrate, let's take an example from anatomy. Bones of sufficient strength to support the bodies of land animals against the downward pull of gravity evolved because the force of gravity has been present and stable over countless generations. To take a behavioral example, humans and many other animals evolved neural circuits for thirst as a motivating force causing them to seek out and consume water. However, natural selection could not have created the neural mechanisms for thirst if cellular need for water had not been consistently present over countless generations.

The general principle is this: natural selection can create genetic adaptations only to situations, adaptive problems, or other conditions of the environment that are regularly present generation after generation, because natural selection requires stable selection criteria over long periods of time to evolve complex adaptations. But in the case of behavioral/psychological adaptations, genetics can only give rather general direction to behavior because the environmental situations or conditions that led to the genetic evolution of the adaptation are themselves rather general categories of events; furthermore, these events are variable in their details but constant in their more abstract common features (Koenigshofer, 2017).

For example, the psychological adaptation, thirst, can guide and motivate the search for water and can motivate its consumption when found, but natural selection cannot code the location of water because the location of water in the environment is too variable from individual to individual and over time, and is therefore too unstable for natural selection to genetically encode the location of water in the environment. This is where learning comes in. Thirst, an innate adaptation, motivates the search for water, but once it is found, the specifics of the location and the quality of the water source must be learned and remembered. In this way, learning supplements genetic information. Learning fills in the detailed information needed to solve a particular adaptive problem or exploit a particular environmental opportunity to adaptive advantage within the specific environment of the individual animal. Genetic information is widespread or even universal throughout a species, whereas learned information is often unique, at least in its details, to the individual.

Genetic information acquired over generations of natural selection is more general and categorical (i.e. more abstract)--e.g. the feeling of thirst means "find and consume water," but doesn't provide the critical information about details such as the location of water. Thus, there arises a general principle: we can expect that whenever information acquired through evolution by natural selection is insufficient to specify a solution to an adaptive problem because more detailed and specific or particular information is needed, then learning mechanisms will evolve that are specific to the problem category (Koenigshofer, 2017).

For example, food selection is important in omnivores such as rats, coyotes, and humans (because they eat a wide variety of potential foods; unlike Koala bears that only eat eucalyptus leaves). In omnivores a specialized form of learning, called taste aversion learning, has evolved. Research shows that learned associations between taste and gastrointestinal illness are readily formed by omnivores, including us, but not in other species such as baleen whales that consume only one kind of food. Have you ever eaten some particular food and then gotten sick later and now you can't stand that food or taste? Genetic evolution equipped omnivores like us with genetic information that allows us to quickly learn taste-illness associations (but not associations between the sight of the food and illness; this is an example of a "biological (i.e. genetic) constraint" on learning). Genetic evolution biologically prepared us to learn taste-illness associations, but it could not tell us which specific tastes we should associate with illness. That detail is left to learning, specific to the experience of the individual during the individual's lifetime.

Another example is that genetic information acquired by natural selection predisposes us, as a species, to learn cause-effect relationships in the environment (see Chapter 14 on Intelligence and Cognition) but doesn't tell us which specific things in one's own particular environment are causally related--again, that detail is left to learning and the learning is guided by genetically evolved predispositions to search for and learn causal relations in the environment (Koenigshofer, 2017).

 

Figure \(\PageIndex{6}\): Web construction by spiders is a complex set of movements directed by circuits in the spider nervous system constructed by information in the spider's DNA. The spider's behavior is so stereotyped that experts who study spiders can identify the species of spider from the structure of its web alone, even if the spider is not present. No learned information is required by the spider to construct its species-typical web. By contrast, human construction of structures is highly dependent upon learned information that has been culturally transmitted across generations. The human brain evolved mechanisms for learning, cultural transmission, comprehension of three-dimensional space, and the ability to visualize in imagination forms like the one depicted here. (Images from Wikipedia Commons; Source of image of spider and web: File:Spider weaving it's web.jpg; https://commons.wikimedia.org/wiki/F...it%27s_web.jpg; by Varun V Vasista; licensed under the Creative Commons Attribution-Share Alike 4.0 International license. Source of image of palace in Spain: File:Dawn Charles V Palace Alhambra Granada Andalusia Spain.jpg; https://commons.wikimedia.org/wiki/F...usia_Spain.jpg; by Jebulon; made available under the Creative Commons CC0 1.0 Universal Public Domain Dedication license).

Slow vs. Fast Mechanisms of Behavioral Change

Genetically organized behavioral adaptations (such as web-building in spiders), like anatomical and physiological adaptations, are organized by genetic information and therefore they change (evolve) only relatively slowly, over many generations by natural selection and other mechanisms of evolution (see Chapter 3). However, for many genes in the organism's phenotype, the gene's expression can be affected by multiple environmental factors. The study of this kind of gene-environment interaction is called epigenetics (see section on epigenetics in Chapter 3).

By contrast, learned behavioral adaptations ("adaptive adjustments") can change moment to moment and, as discussed above, may be transmitted (in some species) to future generations by cultural transmission to the benefit of those future generations. For example, the agricultural practices--learned behavioral adaptations or "adjustments"--upon which we have come to depend for our food supply, were invented by people who died generations ago. However, the death of those who invented these behaviors did not result in the loss of the successful behavioral adaptations upon which we depend for our food supply. Instead, these learned behavioral adaptations have been passed on, and refined, over many generations, to our current generation to its adaptive advantage--not by genetic transmission, but by cultural transmission. Our great capacity for cultural transmission is the single thing that makes us most different from other species, and accounts more than anything else for what we think of as being most characteristically human---technology, art, agriculture, governments, economies, medicine, and science. All would impossible were we, humans, not so powerfully equipped by our brain evolution for efficient cultural transmission of learned behavioral adaptations across generations. By this means, the learning of prior generations is not lost, but remains over generations for each new generation to build upon. Just like flying is a specialty of birds, or swimming is a specialty of fish, cultural transmission is the specialty of the human species.

Summary

In short, learning describes processes whereby information is acquired during the lifetime of an individual animal. Learned information, along with genetic information, helps organize the animal's behavior into adaptive patterns, especially in response to short-term environmental changes where specific, frequently changing details are adaptively significant and therefore must be captured by the organism and put to adaptive use. For example, learning and remembering where a temporary water hole is located on the open savannah is essential to survival for innumerable species that live on the African plains. Because many short-term event details do not regularly recur over generations, such non-recurrent environmental details cannot drive natural selection for genetically evolved instinctual or reflexive adaptations. Instead adaptation to such novel idiosyncratic event details favors the evolution of learning mechanisms (Koenigshofer, 2017). Usually this learned information supplements hereditary (genetic) sources of information in the organization of successful behavioral adaptations--behavioral solutions to the problems of survival and reproduction. As noted above, behavior is one way that organisms (at least animals) adapt. Behavior becomes organized into adaptive patterns by information (i.e. behavior is not random, but guided by laws of nature which govern chances of survival and reproduction--laws of evolution and the laws of learning). The information that organizes behavior comes from the genes after having been perfected by eons of genetic evolution by natural selection. That organizing information can also come from learning by an individual animal during its lifetime (laws which govern learning are also organized to enhance survival and reproduction--see sections below).

Adaptive behaviors can be transmitted to future generations. If the behavior is organized by information in the genes, then that behavior can be transmitted by genetic transmission (heredity). If the information for a particular behavior comes from learning (and is stored in memory), then the behavior can be transmitted to future generations, not by genetic transmission, but instead by cultural transmission (in some species, as noted above). Thus behavioral adaptations in species capable of cultural transmission can undergo not only genetic evolution (true for behaviors organized by genetic information contained in DNA, and also true of anatomical and physiological adaptations) but also cultural evolution (examples of cultural evolution are the development of human technology, modern medical practices, agricultural practices, economies, science, governmental systems and so on, leading over time to generally improved human adaptation). Cultural transmission and the resulting cultural evolution gives our species great survival advantage. Cultural transmission accounts for the success of the human species more than any other single factor.

References

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Attributions

Section 10.1, "Learning, Genes, and Adaptation" is original material written by Kenneth A. Koenigshofer, PhD. and is licensed under CC BY 4.0.

Images from Wikimedia Commons.