A learned behavior is a behavior that is not natural but rather developed through socialization.

Most behaviors, besides basic animalistic Trial and Error Learning

instincts, are learned from somewhere or someone. Trial and error learning is learning from multiple attempts. Here to seek out an answer multiple attempts are made after the primary attempt if
Habituation an answer isn’t found, another attempt is being made, and still not found solution another attempt is additionally being made, then on until
Habituation is a simple form of learning in which an animal stops responding to a stimulus after a period of repeated exposure. This is a form unless an answer is found.
of non-associative learning, as the stimulus is not associated with any punishment or reward. Prairie dogs typically sound an alarm call when In other words, consistent with the trial and error method learning is that the stamping of correct responses and stamping out incorrect
threatened by a predator, but they become habituated to the sound of human footsteps when no harm is associated with this sound, responses.
therefore, they no longer respond to them with an alarm call. In this example, habituation is specific to the sound of human footsteps, as the Edward Lee Thorndike, an American psychologist, is that the first promoter of Trial and Error learning. it’s also called Thorndike’s trial and
animals still respond to the sounds of potential predators. error theory of learning. he’s known for his works on animal psychology and academic psychology. He studied learning this theory employing
a cat through the “puzzle box.”
Imprinting This theory falls under the category of behavioristic theories and follows the stimulus-response theory. Thorndike says “Learning is
In psychology and ethology, Imprinting is a type of learning that occurs at a particular age or a life stage that is rapid and independent of the connecting, the mind is man’s connection system”. Learning is caused by the formation of a connection between stimuli and response.
species involved. It was first used to describe situations in which an animal or person learns the characteristics of some stimulus, which is This theory of learning usually takes place in trial and error. As its name suggests, to seek out an answer we usually need to undergo trials
therefore said to be "imprinted" onto the subject. Imprinting is hypothesized to have a critical period. Hatchling ducks recognize the first and errors. to urge to the answer we made the amount random of trials because the number of trials increases errors are decreased. the
adult they see, their mother, and make a bond with her. A familiar sight is ducklings walking or swimming after their mothers. This is another most idea of trial and error learning is “several trials end in decreased errors.”
type of non-associative learning, but is very important in the maturation process of these animals as it encourages them to stay near their Latent Learning
mother so they will be protected, greatly increasing their chances of survival. However, if newborn ducks see a human before they see their The word latent means hidden or underlying, i.e., something that is present but not always obvious. But what is latent learning? Latent
mother, they will imprint on the human and follow it in just the same manner as they would follow their real mother. learning is learning that only becomes apparent after an incentive is introduced. For example, a teenager riding in a car with a parent takes
Signs of Imprinting note of how the parent is driving; when to use a signal, when to yield, etc. But the teenager does not display this knowledge until the
incentive of a driver license is obtainable. Latent learning describes learning that is hidden or not observed until a reason exists or motivation
There are a few signs of imprinting that can determine if a newborn animal has made an attachment. The main determination in Lorenz's exists to use it.
research was tested by separating the baby animal away from its mother, the object, or animal the young assumably imprinted on. After they A lot of information is learned during a lifetime but not everything has an immediate use. Sometimes knowledge is filed away until a situation
have been separated for an extended amount of time, the object or animal was reintroduced for the newborn to follow. If the baby followed arises where it would be useful. This was what psychologists termed behaviorism. Learning was only validated once a task could be
the theorized object or animal, it would have most likely imprinted on it. performed or a behavior was displayed to prove something was learned-but only in order to receive something desirable. Behaviorism is the
Types of Imprinting school of though that since learning could not be seen, only actions or behaviors that could be seen were proof that learning took place, and
There are three primary types of imprinting that have been discovered in psychological research. that reinforcements were needed to compel learning. Learning how to cook is a behavior that is displayed so that a person can eat. Learning
1. Filial imprinting 2. Sexual imprinting 3.Limbic imprinting a new language is a behavior that is displayed so a person can communicate with others. Learning a new skill is a behavior a person
1. Filial Imprinting displays in order to keep a job and make a living.
What Is Insight Learning?
Filial imprinting is the most widely known type of imprinting that influences the connection between young and their parents or siblings. This
Insight learning is a process that leads to a sudden realization regarding a problem. Often, the learner has tried to understand the
type of behavioral bond allows the young to visually imprint on their parents as a means of survival. This unique bond also ensures that the
problem, but steps away before the change in perception occurs. Insight learning is often compared to trial-and-error learning, but it’s
young will recognize their parent or even different members within their species which is useful for reproduction.
slightly different.
2. Sexual Imprinting
Rather than just trying different random solutions, insight learning requires more comprehension. Learners aim to understand the
Sexual imprinting is a type of behavioral trait that affects an animal's inclination towards a sexual mate from learned behavior, usually very relationships between the pieces of the puzzle. They use patterns, organization, and past knowledge to solve the problem at hand.
early in life. This type of imprinting behavior influences mate selection later in life. It can be altered to make an animal prone to Is Insight Learning Only Observed In Humans?
characteristics or choose against specific features. Humans aren’t the only species that learn with insight. Not all species use this process – just the ones that are closest to us intellectually.
The Westermarck effect is a type of aversion imprinting that may help select against breeding. This effect is also known Insight learning was first discovered not by observing humans, but by observing chimps.
as reverse sexual imprinting and theorizes that people who live together, usually in a family unit, early in life will most likely not develop In the early 1900s, Wolfgang Köhler observed chimpanzees as they solved problems. Köhler’s most famous subject was a chimp named
sexual feelings for each other. Sultan. The psychologist gave Sultan two sticks of different sizes and placed a banana outside of Sultan’s cage. He watched as Sultan
3. Limbic imprinting looked at the sticks and tried to reach for the banana with no success. Eventually, Sultan gave up and got distracted. But it was during this
Some suggest that prenatal, perinatal and post-natal experiences leave imprints upon the limbic system, causing lifelong effects and this time that Köhler noticed Sultan having an “epiphany.” The chimp went back to the sticks, placed one inside of the other, and used this to
process is identified as limbic imprinting. The term is also described as the human emotional map, deep-seated beliefs, and values that are bring the banana to him.
stored in the brain's limbic system and govern people's lives at the subconscious level. Four Stages of Insight Learning
Multiple models have been developed, but the four-stage model is the most popular. The four stages of insight learning are preparation,
Conditioned Behavior incubation, insight, and verification.
Conditioned behaviors are types of associative learning, where a stimulus becomes associated with a consequence. During operant Preparation
conditioning, the behavioral response is modified by its consequences, with regards to its form, strength, or frequency. The process begins as you try to solve the problem. You have the materials and information in front of you and
Classical Conditioning begin to make connections. Although you see the relationships between the materials, things just haven’t
In classical conditioning, a response called the conditioned response is associated with a stimulus that it had previously not been associated “clicked” yet. This is the stage where you start to get frustrated.
with, the conditioned stimulus. The response to the original, unconditioned stimulus is called the unconditioned response. The most cited Incubation
example of classical conditioning is Ivan Pavlov’s experiments with dogs (Figure 2). In Pavlov’s experiments, the unconditioned response During the incubation period, you “give up” for a short period of time. Although you’ve abandoned the project,
was the salivation of dogs in response to the unconditioned stimulus of seeing or smelling their food. The conditioning stimulus that your brain is still making connections on an unconscious level.
researchers associated with the unconditioned response was the ringing of a bell. During conditioning, every time the animal was given Insight
food, the bell was rung. This was repeated during several trials. After some time, the dog learned to associate the ringing of the bell with When the right connections have been made in your mind, the “a-ha” moment occurs. Eureka! You have an
food and to respond by salivating. After the conditioning period was finished, the dog would respond by salivating when the bell was rung, epiphany!
even when the unconditioned stimulus, the food, was absent. Thus, the ringing of the bell became the conditioned stimulus and the Verification
salivation became the conditioned response. Now, you just have to make sure that your epiphany is right. You test out your solution and hopefully, it works! This is a great moment in your
Operant Conditioning learning journey. The connections you make solving this problem are likely to help you in the future.
In operant conditioning, the conditioned behavior is gradually modified by its consequences as the animal responds to the stimulus. A
major proponent of such conditioning was psychologist B.F. Skinner, the inventor of the Skinner box. Skinner put rats in his boxes that Reasoning: Meaning, Definition and Types
contained a lever that would dispense food to the rat when depressed. While initially the rat would push the lever a few times by accident, it It is one of the best forms of controlled thinking consciously towards the solution of a problem. It is realistic in the sense that the solution is
eventually associated pushing the lever with getting the food. This type of learning is an example of operant conditioning. Operant learning is sought always in reference to the reality of the situation. We can solve many problems in our day-dreams, dreams and imaginations but they
the basis of most animal training. The conditioned behavior is continually modified by positive or negative reinforcement, often a reward are unrealistic solutions.
such as food or some type of punishment, respectively. In this way, the animal is conditioned to associate a type of behavior with the As Sherman defined, “reasoning is a process of thinking during which the individual is aware of a problem identifies,
punishment or reward, and, over time, can be induced to perform behaviors that they would not have done in the wild, such as the “tricks” evaluates, and decides upon a solution”.
dolphins perform at marine amusement park shows Reasoning is used not only when we want to solve an immediate problem but also when we anticipate future problems.
Reasoning plays a significant role in one’s adjustment to the environment. It not only determines one’s cognitive activities but also influences
Cognitive Learning
the behaviour and personality.
Classical and operant conditioning are inefficient ways for humans and other intelligent animals to learn. Some primates, including humans,
Definitions of Reasoning:
are able to learn by imitating the behavior of others and by taking instructions. The development of complex language by humans has made
1. “Reasoning is a stepwise thinking with a purpose or goal in mind” —Garrett.
cognitive learning, the manipulation of information using the mind, the most prominent method of human learning. In fact, that is how
2. “Reasoning is the term applied to highly purposeful, controlled and selective thinking”—Gates.
students are learning right now by reading this book. As students read, they can make mental images of objects or organisms and imagine
3. “Reasoning is the word used to describe the mental recognition of cause and effect relationships, it may be the prediction of an event from
changes to them, or behaviors by them, and anticipate the consequences. In addition to visual processing, cognitive learning is also
an observed cause or the inference of a cause from an observed event”—Skinner.
enhanced by remembering past experiences, touching physical objects, hearing sounds, tasting food, and a variety of other sensory-based
Thus reasoning is a highly specialized thinking which helps an individual to explore mentally the cause and effect relationship of an event or
inputs. Cognitive learning is so powerful that it can be used to understand conditioning in detail. In the reverse scenario, conditioning cannot
solution of a problem by adopting some well-organized systematic steps based on previous experience combined with present observation.
help someone learn about cognition.
Types of Reasoning:
Reasoning may be classified into two types.

1. Inductive reasoning:
It is a specialized thinking aimed at the discovery or construction of a generalized principle by making use of particular cases, special A bee-hive has elaborate homeostatic mechanisms that maintain a constant temperature, water
examples and identifying of elements or relations. balance and nutrient level. When the bee-hive is hot, worker bees fan air throughout it and cool
2. Deductive reasoning: it off. When the hive is short of water, then the nurse workers turn to the nearest workers and
It is the ability to draw some logical conclusions from known statement or evidences. Here one starts with already known or established signal a need for water. The latter then gives what water it has and then turns to its neighbours.
generalized statement or principle and applies it to specific cases. For example, all human beings are mortal you are a human being, Thus, the message of the shortage of water moves from bee to bee until it reaches a worker who
therefore, you are mortal. leaves the hive and returns with more water. When a predator or parasite enters the hive, worker
Henry has categorized three types of deductive reasoning: bees rush forward and defend the colony and in the process they may die. Thus, a bee-hive works
i. Conditioned reasoning: as a marvelous society.
It is the reasoning tied down by some specific condition such as the following.
ii. Categorical reasoning: Shoaling and schooling
This type of reasoning is based on some categorical statements.
iii. Linear reasoning: Any group of fish that stay together for social reasons are shoaling, and if the group is
This type of reasoning involves straight forward relationships among elements. swimming in the same direction in a coordinated manner, they are schooling. In common
usage, the terms are sometimes used rather loosely. About one quarter of fish species shoal all
Social Behaviour of Honeybees their lives, and about one half shoal for part of their lives.
Honeybees have developed one of the most highly organised societies. They live in colonies. The colonies of the honeybees are perennial and Fish derive many benefits from shoaling behaviour including defence against predators (through
a good colony of honeybee may consist of 50,000 to 80,000 individuals. It comprises of one fertile female or the queen, a few hundreds of better predator detection and by diluting the chance of individual capture), enhanced foraging
male bees or drones and the rest are sterile females or workers. success, and higher success in finding a mate. It is also likely that fish benefit from shoal membership through increased hydrodynamic
The queen lays the eggs which develops into new workers, drones and queens. She also emits a complex series of chemical secretions, the efficiency.
pheromones that regulate much of the behaviour of the workers. Both queen and worker are genetically diploid. Queens, however, are fed a Fish use many traits to choose shoalmates. Generally they prefer larger shoals, shoalmates of their own species, shoalmates similar in size
special rich larval food; a white, foamy, yogurt-like royal jelly. and appearance to themselves, healthy fish, and kin (when recognized).
It is necessary for normal queen size and sexual development. Workers too, can develop into sexually reproducing females, but their An aggregation of fish is the general term for any collection of fish that have gathered together in some locality. Fish aggregations can be
reproductive organs are kept undeveloped through the effect of the queen’s pheromones. Drones are genetically haploid and are produced by structured or unstructured. An unstructured aggregation might be a group of mixed species and sizes that have gathered randomly near
the laying of unfertilized eggs by parthenogenesis. Drones are generally produced at the same time as the new queen. some local resource, such as food or nesting sites.
A. Worker: If, in addition, the aggregation comes together in an interactive, social way, they may be said to be shoaling. Although shoaling fish can
The worker honeybee (Fig. 1) is the smallest member of the colony. It is black or brown in colour and its mouth parts are of the rasping and relate to each other in a loose way, with each fish swimming and foraging somewhat independently, they are nonetheless aware of the other
lapping type, to facilitate the collection of nectar and pollen. The legs are covered with hair and are adapted for gathering pollen. members of the group as shown by the way they adjust behaviour such as swimming, so as to remain close to the other fish in the group.
The mesothoracic legs bear a pollen brush and a spine-like pollen spur to remove pollen from pollen baskets. The metathoracic legs has a Shoaling groups can include fish of disparate sizes and can include mixed-species subgroups.
depression called the pollen basket. The last four visible segments of the abdomen have modified cells on the ventral surface that secretes If the shoal becomes more tightly organised, with the fish synchronising their swimming so they all move at the same speed and in the same
wax, which is essential for making the hive. direction, then the fish may be said to be schooling. Schooling fish are usually of the same species and the same age/size. Fish schools
A worker honeybee lives for about 6 weeks as an adult and her activities are to some extent synchronized with her physiology. She spends the move with the individual members precisely spaced from each other. The schools undertake complicated manoeuvres, as though the
first three days in cleaning the cells and then begins feeding the older larvae a mixture of pollen and honey. schools have minds of their own.
This she picks up from the storage cells of the hive. The workers then start secreting the so-called royal jelly, and from about the 6th to the Social interaction
14th day of her life she feeds this secretion to the younger larvae and any queen larvae in the hive. Support for the social and genetic function of aggregations, especially those formed by fish, can be seen in several aspects of their
For a brief period, the royal jelly is fed to all larvae, but those that will become the queen are fed with royal jelly throughout their larval behaviour. For instance, experiments have shown that individual fish removed from a school will have a higher respiratory rate than those
period. Form the 10th day onwards, the worker’s wax-secreting glands on the abdomen become active and, at the same time, the pharyngeal found in the school. This effect has been attributed to stress, and the effect of being with conspecifics therefore appears to be a calming one
glands begin to regress. The workers then gradually change their behaviour from feeding larvae to cell construction. and a powerful social motivation for remaining in an aggregation.
About the 18th day, the worker may leave the colony occasionally for a few brief orientation flights. At this stage, she may be found guarding Foraging advantages
the hive entrance and inspecting incoming bees. From the 21st day onwards the It has also been proposed that swimming in groups enhances foraging success. This ability was demonstrated by Pitcher and others in their
workers change into a forager, bringing back nectar, pollen and water, and study of foraging behaviour in shoaling cyprinids. In this study, the time it took for groups of minnows and goldfish to find a patch of food was
continue this function for the rest part of its life. quantified. The number of fishes in the groups was varied, and a statistically significant decrease in the amount of time necessary for larger
B. Drone: groups to find food was established. Further support for an enhanced foraging capability of schools is seen in the structure of schools of
Drones are intermediate in size (Fig. 5.29) but considerably stouter and broader. predatory fish. Partridge and others analysed the school structure of Atlantic bluefin tuna from aerial photographs and found that the school
They possess large eyes, small pointed mandibles and lack wax-producing assumed a parabolic shape, a fact that was suggestive of cooperative hunting in this species.
glands, pollen-collecting apparatus and a sting. The drones do not do any work Reproductive advantages
and, if not fed by the workers, they die. They exist only to mate with the queen. A third proposed benefit of fish groups is that they serve a reproductive function. They provide increased access to potential mates, since
C. Queen: finding a mate in a shoal does not take much energy. And for migrating fish that navigate long distances to spawn, it is likely that the
The queen being the only fertile female of the hive, have immensely developed navigation of the shoal, with an input from all the shoal members, will be better than that taken by an individual fish. Forage fish often make
ovaries. The queen is elongated with a long tapering abdomen with great migrations between their spawning, feeding and nursery grounds. Schools of a particular stock usually travel in a triangle between
comparatively shorter wings and legs. She has pointed mandibles and shorter these grounds.
mouth parts and sting with no barbs. Hydrodynamic efficiency
She alone lays eggs and is the mother of almost all the members of the hive. She lives for several successive years laying about 1-200 eggs a This theory states that groups of fish may save energy when swimming together, much in the way that bicyclists may draft one another in a
day and laying about 15 lakh eggs during her life time. When a hive of honeybees prepare to swarm or when an old queen becomes weak, the peloton. Geese flying in a Vee formation are also thought to save energy by flying in the updraft of the wingtip vortex generated by the
regulating pheromones of the queen also become weak. This serves as a signal for workers to begin raising a new queen. previous animal in the formation. Increased efficiencies in swimming in groups have been proposed for schools of fish and Antarctic krill.
Swarming: Predator avoidance
Swarming takes place during the spring or early summer. This relieves over-crowding and also provides a means of founding new colonies.
One potential method by which fish schools might thwart predators is the "predator confusion effect" proposed and demonstrated by Milinski
Prior to a swarm, a large number of queen and drone cells are constructed. Then on a clear day the old queen leaves the hive with a group of
and Heller (1978). This theory is based on the idea that it becomes difficult for predators to choose individual prey from groups because the
old workers and drones.
many moving targets create a sensory overload of the predator's visual channel. Milinski and Heller's findings have been corroborated both
Left behind in the hive are the young workers and several new queens who are still in their cells but approaching the time of emergence. The
in experiment and computer simulations.] "Shoaling fish are the same size and silvery, so it is difficult for a visually oriented predator to pick
first queen that hatches out becomes the mistress of the parent colony. This new queen kills any other newly-hatched rivals.
an individual out of a mass of twisting, flashing fish and then have enough time to grab its prey before it disappears into the shoal."
After about a week of her emergence, the queen flies off to mate. By this time, drones also have left the hive and have aggregated in
traditional sites in large flying clouds. The drones from many neighbouring hives may also combine in such a cloud called drone cloud. When How fish school
the new queen approaches such a drone cloud, the drones rush to the queen and several of them may mate with a queen in succession. Fish schools swim in disciplined phalanxes, with some species, such as herrings, able to stream up and down at impressive speeds, twisting
Mating occurs in mid-air. The sperms are stored in a spermatheca or sperm-reservoir of the queen, which fertilizes her eggs as long as she this way and that, and making startling changes in the shape of the school, without collisions. It is as if their motions are choreographed,
lives. The male dies after mating, with its genital parts remaining in the abdomen of the queen. It is later removed by the queen or by though they are not. There must be very fast response systems to allow the fish to do this. Young fish practice schooling techniques in pairs,
workers. The queen, after mating, returns back to the hive not to leave it again, until she grows old and leads a prime swarm. and then in larger groups as their techniques and senses mature. The schooling behaviour develops instinctively and is not learnt from older
Bee-hive: fish. To school the way they do, fish require sensory systems which can respond with great speed to small changes in their position relative
The bee-hive consists of two layers of hexagonal chambers or cells made by the bees wax secreted from the abdomen of the worker bees. to their neighbour. Most schools lose their schooling abilities after dark, and just shoal. This indicates that vision is important to schooling.
Before the wax is used it is masticated and mixed with secretions of cephalic glands to convert it into a plastic substance. The importance of vision is also indicated by the behaviour of fish who have been temporarily blinded. Schooling species have eyes on the
The bee-hive consists of two types of cells: sides of their heads, which means they can easily see their neighbours. Also, schooling species often have "schooling marks" on their
1. Storage cells which contains honey and pollen, usually built near the top. shoulders or the base of their tails, or visually prominent stripes, which provide reference marks when schooling, similar in function to
2. Brood cells generally occupy the lower and central portions. passive markers in artificial motion capture. However fish without these markers will still engage in schooling behaviour though perhaps not
They are of three types (Fig. 2 ) and differ in shape and size: as efficiently.
(i) Worker cells are small like the honey cells. Other senses are also used. Pheromones or sound may also play a part but supporting evidence has not been found so far. The lateral line
(ii) Drone cells are slightly larger. is a line running along each side of the fish from the gill covers to the base of the tail. In laboratory experiments the lateral lines of schooling
(iii) Queen cells are enoromous, cylindrical or vase-shaped in structure. Once used these cells cannot be used again. fish have been removed. They swam closer, leading to a theory that the lateral lines provide additional stimuli input when the fish get too
close. The lateral-line system is very sensitive to changes in water currents and vibration in the water. It uses receptors called neuromasts,
 each of which is composed of a group of hair cells. The hairs are surrounded by a protruding jelly-like cupula, typically 0.1 to 0.2 mm long. ● relieve physical pain
The hair cells in the lateral line are similar to the hair cells inside the vertebrate inner ear, indicating that the lateral line and the inner ear “Altruism is also associated with post-traumatic growth,” says Manly.
share a common origin. 5 ways to foster altruism
Describing shoal structure 1. Consider mindfulness
It is difficult to observe and describe the three dimensional structure of real world fish shoals because of the large number of fish involved. Mindfulness is the practice of staying in the present moment. When you learn to disengage from distractions, this may open you up to notice
Techniques include the use of recent advances in fisheries acoustics. the world around you right here, right now.
Parameters defining a fish shoal include: 2. Try to grow in empathy
● Shoal size – The number of fish in the shoal. A remote sensing technique has been used near the edge of the continental To enhance your empathy, you may want to consider:
shelf off the east coast of North America to take images of fish shoals. The shoals – most likely made up of Atlantic herring, ● trying to make direct eye contact when someone talks with you
scup, hake, and black sea bass – were said to contain “tens of millions” of fish and stretched for “many kilometers”. ● attending gatherings with diverse groups of people
● Density – The density of a fish shoal is the number of fish divided by the volume occupied by the shoal. Density is not ● following people on social media who are different from you
necessarily a constant throughout the group. Fish in schools typically have a density of about one fish per cube of body length. 3. Starting small may help
● Polarity – The group polarity describes the extent to which the fish are all pointing in the same direction. In order to determine “Many people can incorporate altruism into their everyday life with small acts of kindness,” says Myszak.
this parameter, the average orientation of all animals in the group is determined. For each animal, the angular difference 4. Consider a group effort
between its orientation and the group orientation is then found. The group polarity is the average of these differences. If you feel a little nervous about walking into a new or vulnerable situation, you may find it helpful to reach out to a friend or loved one.
● Nearest neighbour distance – The nearest neighbour distance (NND) describes the distance between the centroid of one fish 5. Try to find the right fit
(the focal fish) and the centroid of the fish nearest to the focal fish. This parameter can be found for each fish in an aggregation “As you develop a practice of altruism, try to find a good fit to ensure that you don’t overextend yourself,” says Manly.
and then averaged. Care must be taken to account for the fish located at the edge of a fish aggregation, since these fish have
no neighbour in one direction. The NND is also related to the packing density. For schooling fish the NND is usually between Reproductive strategies and mating systems: the foundations of sexual selection in animals
one-half and one body length. For many years, it has been been thought that living beings were basically organisms that are born, grow, reproduce and
● Nearest neighbour position – In a polar coordinate system, the nearest neighbour position describes the angle and distance die, or that they were organisms that fulfilled their physiologic functions: nutrition, relation and reproduction. Such
of the nearest neighbour to a focal fish. statements are, with some nuances, valid definitions of a “living being” and they all share a common denominator: they
● Packing fraction – The packing fraction is a parameter borrowed from physics to define the organization (or state i.e. solid,
mention the reproductive function as a characteristic. This gives us a hint of the importance of such function for
liquid, or gas) of 3D fish groups. It is an alternative measure to density. In this parameter, the aggregation is idealized as an
individuals. In this article, we are going to discover why sexual selection is so essential.
ensemble of solid spheres, with each fish at the center of a sphere. The packing fraction is defined as the ratio of the total
Most individuals — if not all — of every animal species engage in some type of sexual behaviour throughout their lifetime.
volume occupied by all individual spheres divided by the global volume of the aggregation. Values range from zero to one,
where a small packing fraction represents a dilute system like a gas. The objective of such behaviour in individuals is to pass on their genes to the next generation, ensuring their viability.
● Integrated conditional density – This parameter measures the density at various length scales and therefore describes the However, their success will depend on the different sexual strategies of each species and gender, as well as on each mating
homogeneity of density throughout an animal group. system:
● Pair distribution function – This parameter is usually used in physics to characterize the degree of spatial order in a system Reproductive strategies in males and females
of particles. It also describes the density, but this measure describes the density at a distance away from a given point. The reproductive strategies of males and females are related to the characteristics of their respective gametes: while
Cavagna et al. found that flocks of starlings exhibited more structure than a gas but less than a liquid. sperm cells are abundant, “cheap” and easy to replace, ovules are scarce and more costly to obtain; this leads males and
Altruism: females to carry out different reproductive strategies. Nonetheless, the role of each gender during reproduction is
Altruism is the selfless act of helping others without expecting anything in return. “It is often considered one of the defining characteristics of determined by environmental factors (for example, by the distribution of resources) and vital cycles (if oestrus is
what it means to be human,” says Dr. Jessica Myszak, a psychologist in Glenview, Illinois. synchronised or not).
Types of altruism Generally, males try to copulate as many times as possible and compete over females, while females procure the best
Experts have long been fascinated by the motivations of human cooperation, noting four distinct types of altruism: care for their offspring and choose the most fit males. To do so, they analyse the parental care that the male can provide
1. Kin altruism. Kin altruism involves behaving in such a way as to benefit a genetic relative’s chances of survival or reproduction at and they favour males with attributes that indicate genetic quality.
some cost to one’s own chances. An obvious example of this kind of altruism is parental investment, whereby an individual risks its There is, therefore, a conflict between males and females, as the former try to mate indiscriminately, whereas females act
safety or limits its future fecundity by caring for its offspring. It happens when you unselfishly support your family members and more intentionally. For certain species, the conflict is even more severe: in some insects, for example, the males’ semen
loved ones or make personal sacrifices on their behalf. Ex- caregiving for a relative with a chronic condition contains substances that affect the females’ brain, making them behave in a way that increases their chances to reproduce
● donating blood or a major organ to your sibling successfully, but this substance also shortens the female’s life.
● taking a second job to pay for your child’s education Mating systems
● fostering or adopting children For this conflict between males and females not to be insurmountable, there are several mating options:
Monogamy
2. Reciprocal altruism. In evolutionary biology, reciprocal altruism is a behaviour whereby an organism acts in a manner that A system through which a couple establishes an exclusive sexual relationship, particularly during reproductive cycles, that
temporarily reduces its fitness while increasing another organism's fitness, with the expectation that the other organism will act in a
can last an entire lifetime. This system is predominant amongst birds (over 90%), although some intercourse with other
similar manner at a later time.
mates is also common. Monogamy occurs when:
The concept was initially developed by Robert Trivers to explain the evolution of cooperation as instances of mutually altruistic acts. ● Males are unable to monopolise females or resources, or when mating with several females has no reproductive
The concept is close to the strategy of "tit for tat" used in game theory. benefits.
It occurs when you help someone knowing that, at some point, they may help you in the future as well. ex- ● The likelihood of successful breeding is higher with the help of both parents, and males can contribute to the
● loaning a friend money, who helped you cover bills raising of the offspring.
● helping your classmate with a project, who helped you study Polygyny
● holding the elevator for your colleague, who brought you a coffee A system in which a male monopolises harem of females and mates with them on a regular basis. The females have an
exclusive sexual relationship with said male. This is the most common system amongst mammals (over 80%) and
3. Cultural group altruism. It involves supporting someone who’s part of a group you are associated with, including ethnic and social uncommon amongst birds (barely 2%). It occurs when males are able to monopolise females (no oestrus synchronisation)
groups. ex- and/or resources.
● starting a nonprofit for a cause you care about Polyandry
● donating money to an organization that supports people of your background In this case, it is the female who establishes exclusive mating relationships with several males. Polyandry is a common
● donating items to people in your religious group mating system amongst fish, as well as amongst some males and birds. It occurs when:
● Food is scarce, or in areas with an abundance of food, but that have been affected by a natural disasters.
4. Pure altruism. It involves helping someone from a place of empathy knowing you will see no benefit, often in high stake situations. ● Males assume the care of their offspring, or when the male ratio is a lot higher.
ex- Promiscuity
● holding open the door for a stranger
Both sexes mate randomly with different individuals. This system occurs in many pelagic fish, and when there are large
● helping an older adult walking with a cane cross the street
amphibian populations in small ponds. It also occurs amongst some birds, but to a lesser extent (barely 6% of bird species).
● volunteering at a soup kitchen
Female mate choice
● bringing a lost animal to the shelter
Altruism and prosocial behavior may: For mating to take place, individuals have to pair up first. Females take several factors into account in order to choose a
● enhance social connections mate:
● increase your sense of community ● Hunt for natural resources. The likes for a male to be chosen by a female are higher when it defends a territory
● improve your mood with abundant resources (water, vegetation, etc.). This type of choice occurs, for example, in frogs.
● protect against cognitive decline ● Good genes. Females tend to choose males that are easy to spot. This is why males use bright colours, ornaments
● Trusted Source and extrinsic elements during courtship. The flame bowerbird (Sericulus aureus) decorates the nest it builds during
● provide a sense of purpose courtship with a variety of gifts (fruits, feathers, flowers…), increasing its likelihood of being selected.
● reduce a sense of loneliness Consequences of sexual selection

Due to the different strategies and factors explained in this article, sexual selection has a series of consequences for
Genetic heritability cost of sex
species and individuals:
A sexually reproducing organism only passes on ~50% of its own genetic material to each L2 offspring. This is a consequence of the fact
● Sexual dimorphism, meaning variations in external physiognomy amongst males and females of the same species.
that gametes from sexually reproducing species are haploid. Again, however, this is not applicable to all sexual organisms
It tends to be more pronounced in polygamous species.
DNA repair and complementation
● (Apparent) conflict with natural selection. The high reproductive success of males with exaggerated features
The repair and complementation hypothesis assumes that genetic recombination is fundamentally a DNA repair process, and that when it
compensates for other costs: for example, having bright colours may be an advantage in sexual selection, but it can
occurs during meiosis it is an adaptation for repairing the genomic DNA which is passed on to progeny. Recombinational repair is the only
also cause a higher predation rate. repair process known which can accurately remove double-strand damages in DNA, and such damages are both common in nature and
● Conflict between growing and maturing. Specifically amongst polygynyc males, there is intrasexual competition ordinarily lethal if not repaired. For instance, double-strand breaks in DNA occur about 50 times per cell cycle in human cells (see naturally
(individuals of the same sex compete over the opposite sex), which leads to a later sexual maturity that females, as occurring DNA damage). Recombinational repair is prevalent from the simplest viruses to the most complex multicellular eukaryotes. It is
well as a shorter reproductive period. effective against many different types of genomic damage, and in particular is highly efficient at overcoming double-strand damages. Studies
● Differences in reproductive potential. Such differences are very significant amongst polygynyc males, and can of the mechanism of meiotic recombination indicate that meiosis is an adaptation for repairing DNA. These considerations form the basis for
considerably increase if their behaviour entails high risk. For example, a deer can increase its reproductive potential the first part of the repair and complementation hypothesis.
if it fights and competes with other males over a harem of females, but this also increases their mortality rate. Deleterious mutation clearance
Mutations can have many different effects upon an organism. It is generally believed that the majority of non-neutral mutations are
Evolution of sexual reproduction deleterious, which means that they will cause a decrease in the organism's overall fitness.
Sexual reproduction is an adaptive feature which is common to almost all multicellular organisms and various unicellular organisms, with
some organisms being incapable of asexual reproduction. Currently the adaptive advantage of sexual reproduction is widely regarded as a Evading harmful mutation build-up
major unsolved problem in biology. As discussed below, one prominent theory is that sex evolved as an efficient mechanism for producing While DNA is able to recombine to modify alleles, DNA is also susceptible to mutations within the sequence that can affect an organism in a
variation, and this had the advantage of enabling organisms to adapt to changing environments. Another prominent theory, also discussed negative manner. Asexual organisms do not have the ability to recombine their genetic information to form new and differing alleles. Once a
below, is that a primary advantage of outcrossing sex is the masking of the expression of deleterious mutations.[1] Additional theories mutation occurs in the DNA or other genetic carrying sequence, there is no way for the mutation to be removed from the population until
concerning the adaptive advantage of sex are also discussed below. Sex does, however, come with a cost. In reproducing asexually, no time another mutation occurs that ultimately deletes the primary mutation. This is rare among organisms.
nor energy needs to be expended in choosing a mate. And if the environment has not changed, then there may be little reason for variation, Removal of deleterious genes
as the organism may already be well adapted. Sex also halves the amount of offspring a given population is able to produce. Sex, however, It assumes that the majority of deleterious mutations are only slightly deleterious, and affect the individual such that the introduction of each
has evolved as the most prolific means of species branching into the tree of life. Diversification into the phylogenetic tree happens much additional mutation has an increasingly large effect on the fitness of the organism. This relationship between number of mutations and
more rapidly via sexual reproduction than it does by way of asexual reproduction. fitness is known as synergistic epistasis.
Evolution of sexual reproduction describes how sexually reproducing animals, plants, fungi and protists could have evolved from a common What is Optimal Foraging Theory?
ancestor that was a single-celled eukaryotic species. The optimal foraging theory (OFT) is a model of how animals should forage to maximize fitness. In ecology, fitness is defined as the ability
Since hypotheses for the origin of sex are difficult to verify experimentally (outside of evolutionary computation), most current work has for an organism to successfully reproduce, which is used as a proxy for how well-adapted an animal is to its environment. This is the basis of
focused on the persistence of sexual reproduction over evolutionary time. The maintenance of sexual reproduction (specifically, of its natural selection in that organisms best adapted to their environment can pass on genetic information to the next generation.
dioecious form) by natural selection in a highly competitive world has long been one of the major mysteries of biology, since both other Foraging for food is essential for maintaining physiological functions, but it also requires energy output for locating, handling, killing, and
known mechanisms of reproduction – asexual reproduction and hermaphroditism – possess apparent advantages over it. Asexual consuming prey. The optimal foraging theory predicts behavioral strategies that maximize net energy gain at the lowest energy cost, thus
reproduction can proceed by budding, fission, or spore formation and does not involve the union of gametes, which accordingly results in a maximizing fitness. The main assumption of the optimal foraging theory is that foraging behaviors have been molded by natural selection
much faster rate of reproduction compared to sexual reproduction, where 50% of offspring are males and unable to produce offspring and that foraging behavior is a trait that can be passed on through generations. The optimal foraging theory is a cost-benefit model and
themselves. suggests there is an optimal way to forage to minimize energy output and maximize energy gain. Many factors influence foraging behavior
Advantages of sex and sexual reproduction and optimal foraging has been studied by ecologists for decades. This lesson will discuss assumptions and constraints of the optimal
Reproductive advantages of the asexual forms are in quantity of the progeny, and the advantages of the hermaphrodite forms are in foraging theory and provide examples in nature to explain animal foraging behavior.
maximal diversity. Transition from the hermaphrodite to dioecious state leads to a loss of at least half of the diversity. So, the primary Impact of the optimal foraging theory
challenge is to explain the advantages given by sexual differentiation, i.e. the benefits of two separate sexes compared to hermaphrodites Constraints
rather than to explain benefits of sexual forms (hermaphrodite + dioecious) over asexual ones.
The optimal foraging theory considers these three constraints when determining the cost-benefit relationship:
Advantages due to genetic variation 1. Temporal - the time it takes to acquire food (locating, handling, and consuming prey)
For the advantage due to genetic variation, there are three possible reasons this might happen. First, sexual reproduction can combine the 2. Energetic - the metabolic costs of foraging
effects of two beneficial mutations in the same individual (i.e. sex aids in the spread of advantageous traits). Also, the necessary mutations 3. Cognitive - determining abundance and type of prey available in the environment, related to learning and memory
do not have to have occurred one after another in a single line of descendants. Second, sex acts to bring together currently deleterious Factors influencing constraints
mutations to create severely unfit individuals that are then eliminated from the population
Foraging behavior is influenced by the time and 'energy' necessary to acquire a food source, the 'availability' of the food source, and the
Protection from major genetic mutation presence of predators or competition in the environment.
In contrast to the view that sex promotes genetic variation, Heng, and Gorelick and Heng reviewed evidence that sex actually acts as a ● Prey size: Larger prey may have more nutritional gain, but the process of handling large prey may not be cost effective. This may
constraint on genetic variation. They consider that sex acts as a coarse filter, weeding out major genetic changes, such as chromosomal be impacted by other characteristics such as shell size and thickness (i.e., shellfish, eggs, crustaceans, nuts, or seeds).
rearrangements, but permitting minor variation, such as changes at the nucleotide or gene level (that are often neutral) to pass through the ● Abundance of prey: Animals tend to forage where food sources are most prevalent. When populations of a primary food source
sexual sieve. decrease, predators may have to switch to a less optimal food source (also termed primary and secondary food sources). This may
Novel genotypes be more costly in terms of temporal and metabolic constraints.
Sex could be a method by which novel genotypes are created. Because sex combines genes from two individuals, sexually reproducing ● Predation: Some animals adapt foraging behaviors to avoid the presence of predators. This includes foraging nocturnally or
populations can more easily combine advantageous genes than can asexual populations. If, in a sexual population, two different foraging in habitats that provide cover. When the type of cover changes due to habitat loss, foraging becomes more costly, and
advantageous alleles arise at different loci on a chromosome in different members of the population, a chromosome containing the two animals must adapt to their new environment.
advantageous alleles can be produced within a few generations by recombination. Examples of the optimal foraging theory
Lions
Increased resistance to parasites
When an environment changes, previously neutral or deleterious alleles can become favourable. If the environment changed sufficiently Lions typically hunt in groups at night or in the cooler hours of the day. They are considered opportunistic hunters, meaning they consume a
rapidly (i.e. between generations), these changes in the environment can make sex advantageous for the individual. Such rapid changes in wide range of prey and will hunt based on prey availability. Lions tend to prey on medium-sized ungulates like zebra, buffalo, and warthogs.
environment are caused by the co-evolution between hosts and parasites. However, they have been known to hunt larger animals such as rhinoceros or hippopotamus, although the success rate is often low. The
Disadvantages of sex and sexual reproduction metabolic cost of taking down larger prey is high and sometimes even fatal. However, if they are successful at hunting larger-sized prey, this
The paradox of the existence of sexual reproduction is that though it is ubiquitous in multicellular organisms, there are ostensibly many will often suffice, and the lion can survive without hunting for at least a week.
inherent disadvantages to reproducing sexually when weighed against the relative advantages of alternative forms of reproduction, such as Squirrels
asexual reproduction. Thus, because sexual reproduction abounds in complex multicellular life, there must be some significant benefit(s) to Squirrels nearly exhaust their metabolic energy during the warmer months in order to hoard acorns, nuts, and seeds during the winter
sex and sexual reproduction that compensates for these fundamental disadvantages. months. This is a trade-off in which they expend little to no energy during the winter, surviving off their hoard. The foraging behavior also
changes based on what is available to them during a particular season. For example, the eastern gray squirrel will differentiate between red
Population expansion at the cost of sex and white oak acorns based on availability and taste. Red oak acorns have more tannin, making them more bitter tasting. If both acorns are
Among the most limiting disadvantages to the evolution of sexual reproduction by natural selection is that an asexual population can grow available, the eastern gray squirrel has been known to choose white oak acorns as their primary food source and red oak acorns as the
much more rapidly than a sexual one with each generation. secondary food source.
Selfish cytoplasmic genes
Sexual reproduction implies that chromosomes and alleles segregate and recombine in every generation, but not all genes are transmitted
together to the offspring. There is a chance of spreading mutants that cause unfair transmission at the expense of their non-mutant
colleagues.