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BIOTRANCE

ANIMAL BEHAVIOUR

Ethology – study of Animal behaviour


Behaviors are categorized as follows:
1. Instinct is behavior that is innate, or inherited.
• In mammals, care for offspring by female parents is innate.
2. Fixed action patterns (FAP) are innate behaviors that follow a regular, unvarying pattern. An FAP
is initiated by a specific stimulus. Typically, the behavior is carried out to completion even if the
original intent of the behavior can no longer be fulfilled.
• When a graylag goose sees an egg outside her nest, she will methodically roll the egg back into the
nest with a series of maneuvers using her beak. An egg outside the nest is the stimulus. However, she
will also retrieve any object that resembles her egg, and once the FAP has begun, she will continue
the retrieval motions until she has completed the motions back to the nest. Even if the egg slips away
or is removed, she completes the FAP by returning an “imaginary” egg to the nest.
• Male stickleback fish defend their territory against other males. The red belly of males is the
stimulus for aggressive behavior. However, as ethologist Nikolaas Tinbergen discovered, any object
with a red underside initiates the same aggressive FAP.
3. Imprinting is an innate program for acquiring a specific behavior only if an appropriate stimulus is
experienced during a critical period (a limited time interval during the life of the animal). Once
acquired, the behavior is irreversible.
• Ethologist Konrad Lorenz discovered that, during the first two days of life, graylag goslings will
accept any moving object as their mother. When Lorenz himself was the moving object, he was
accepted as their mother for life. Any object presented after the critical period, including their real
mother, was rejected.
• Salmon hatch in freshwater streams and migrate to the ocean to feed. When they are reproductively
mature, they return to their birthplace to breed, identifying the exact location of the stream. During
early life, they imprinted the odors associated with their birthplace.
4. Associative learning (association) occurs when an animal recognizes (learns) that two or more
events are connected. A form of associative learning called classical conditioning occurs when an
animal performs a behavior in response to a substitute stimulus rather than the normal stimulus.
• Dogs salivate when presented with food. Physiologist Ivan Pavlov found that if a bell were rung just
before dogs were given food, they would, after repeated experiences, salivate in response to the bell
ringing alone. Dogs associated the ring of the bell (the substitute stimulus) with the presentation of
food (the normal stimulus).
5. Trial-and-error learning (or operant conditioning) is another form of associative learning.
It occurs when an animal connects its own behavior with a particular environmental response. If the
response is desirable (positive reinforcement), the animal will repeat the behavior in order to elicit the
same response (for example, to receive a reward). If the response is undesirable (for example,
painful), the animal will avoid the behavior. This is the basis for most animal training by humans.
• Psychologist B. F. Skinner trained rats to push levers to obtain food or avoid painful shocks.
Learning acquired by association can be forgotten or reversed if the performed behavior no longer
elicits the expected response. The loss of an acquired behavior is called extinction.
6. Habituation is a learned behavior that allows the animal to disregard meaningless stimuli.
• Sea anemones pull food into their mouths by withdrawing their tentacles. If the tentacles are
stimulated with nonfood items (a stick, for example), the tentacles will ignore the stimulus after
several futile attempts to capture the “food.”
7. Observational learning occurs when animals copy the behavior of another animal without having
experienced any prior positive reinforcement with the behavior.
• Japanese monkeys usually remove sand from a potato by holding the potato in one hand and
brushing sand away with the other hand. One monkey discovered that she could more easily brush the
sand away if she held the potato in water. Through observational learning, nearly all of the other
monkeys in the troop learned the behavior.
8. Insight occurs when an animal, exposed to a new situation and without any prior relevant
experience, performs a behavior that generates a desirable outcome.
• A chimpanzee will stack boxes so she can climb them, providing her with access to bananas
previously beyond reach. Some behaviors that appear to be learned may actually be innate behaviors
that require maturation.
For example, birds appear to “learn” to fly by trial and error or by observational learning. However, if
birds are raised in isolation, they will fly on their first try if they are physically capable of flying.
Thus, the ability to fly is innate but can occur only after the bird has physically matured. In general,
inherited behaviors and learning capabilities of animals have evolved because they increase
individual fitness. Innate behavior, such as an FAP, improves fitness by providing a successful and
dependable mechanism for the animal to perform in response to an event that, through evolution, has
become expected. By establishing an FAP, a particular challenge need not be resolved repeatedly by
every generation. In contrast, imprinting allows a certain amount of flexibility. If a mother is killed
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ANIMAL BEHAVIOUR
before her chick hatches, the chick will, through imprinting, choose another nearby bird for its mother
(most likely of the same species). Associative learning allows individuals to benefit from exposure to
unexpected (or novel) repeated events. Once they form an association with the event, they can
respond to the next occurrence more efficiently. Habituation allows individuals to ignore repetitive
events which, from experience, they know are inconsequential. As a result, the animals can remain
focused on other, more meaningful events. Observational learning and insight provide a mechanism
to learn new behaviors in response to unexpected events without receiving reinforcement. This
reduces the time required for new behaviors to be acquired.
Instinctual behavior
 One type of instinctual behavior is fixed action patterns, which are behaviors the animal is
compelled to engage in. For instance, some birds will raise the chicks of other birds if the eggs are put
in their nests during nesting season, because caring for an egg is a fixed action pattern. Another
instinctual behavior is imprinting, wherein a baby animal accepts a person, or even an item, as a
surrogate mother. Sexual behavior is also instinctual, bolstered by play, which helps animals learn
courtship and mating skills. Many of these behaviors are dictated by specific body systems, like the
nervous system, which responds to stimuli in the environment.
Learned behavior
 Learned behavior is important both for wild animals, who must learn specific and new ways to
survive, and for domestic animals that we seek to train. Animals can learn to anticipate that an action
will have a predictable outcome through trial and error, such as dog learning to sit for a treat. This is
called operant conditioning. They can also learn that one event precedes another, such as the sound of
a metal food bowl being moved signaling food being served, which is known as associative learning.
Animals also learn a lot through watching others and mimicry. All of these behaviors allow an animal
to adapt to new situations and problems.
Abnormal behavior
 Identifying behavior patterns enables people to determine when animals are behaving abnormally.
These abnormal behaviors might simply be annoying to animal owners; however, in other instances
they may also be dangerous for the animal and others or even threaten their very survival. For
example, inappropriately aggressive dogs, which might be suffering from disease or trauma, are
potentially dangerous to themselves and others. The behavior may be addressed if it is identified as
abnormal and normal behavior is reestablished. More important to species survival are mating and
raising offspring, and in these cases abnormal behavior that leads to failure to mate or care for
offspring can present a threat to the animal's long-term survival.
Territory and territoriality
In ethology the term territory refers to any sociographical area that an animal of a particular species
consistently defends against conspecifics (and, occasionally, animals of other species). Animals that
defend territories in this way are referred to as territorial.
Classic territories
Territorial animals defend areas that contain a nest, den or mating site and sufficient food resources
for themselves and their young. Defense rarely takes the form of overt fights: more usually there is a
highly noticeable display, which may be visual (as in the red breast of the robin), auditory (as in
much bird song, or the calls of gibbons) or olfactory, through the deposit of scent marks. Many
territorial mammals use scent-marking to signal the boundaries of their territories; the marks may be
deposited by urination, by defecation, or by rubbing parts of the bodies that bear specialised scent
glands against the substrate. For example, dogs and other canids scent-mark by urination and
defecation, while cats scent-mark by rubbing their faces and flanks against objects, as well as by the
notoriously persistently smelly spraying of urine by tomcats. Many prosimians use territorial
marking; for example, the Red-bellied Lemur creates territories for groups of two to ten individuals in
the rainforests of eastern Madagascar by scent marking: the male Diademed Sifaka also scent marks
defended territories in some of these same rainforests. The male Western fence lizard defends a
territory by posturing and combat, but less intensely after the mating season.
Invertebrates which show territorality include some ants and bees, and the owl limpet .
Spraying
Spraying (also known as territorial marking) is behavior used by animals to identify their territory.
Most commonly, this is scent marking, accomplished by depositing strong-smelling chemicals such
as urine at prominent locations within the territory. Often the scent contains carrier proteins, such as
the major urinary proteins, to stabilize the odors and maintain them for longer.
Not only does the marking communicate to others of the same species, but it is also noted by prey
species and avoided. For example felids such as leopards and jaguars mark by rubbing themselves
against vegetation. Some prosimians, such as the Red-bellied Lemur, also use scent marking to
establish a territory. Many ungulates, for example the Blue Wildebeest, use scent marking from two
glands, the preorbital gland and a scent gland in the hoof.
Defense
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Territories may be held by an individual, a mated pair, or a group. Territoriality is not a fixed
property of a species: for example, robins defend territories as pairs during the breeding season and as
individuals during the winter, while some nectarivores defend territories only during the mornings
(when plants are richest in nectar). In species that do not form pair bonds, male and female territories
are often independent, in the sense that males defend territories only against other males, and females
only against other females; in this case, if the species is polygynous, one male territory will probably
contain several female territories, while in some polyandrous species such as the Northern Jacana,
this situation is reversed.
Quite often territories that only yield a single resource are defended. For example, European
Blackbirds may defend feeding territories that are distant from their nest sites, and in some species
that form leks, for example the Uganda kob (a grazing antelope), males defend the lek site (which is
used only for mating).
Territoriality is only shown by a minority of species. More commonly, an individual or a group of
animals will have an area that it habitually uses but does not necessarily defend; this is called its
home range. The home ranges of different groups often overlap, and in the overlap areas the groups
will tend to avoid each other rather than seeking to expel each other. Within the home range there
may be a core area that no other individual group uses, but again this is as a result of avoidance
rather than defense.
Behavioural ecologists have argued that food distribution determines whether a species will be
territorial or not. This however, though true as far as it goes, is too narrow a point of view. As
mentioned above, there are several kinds of territoriality; for example, the defence of lek areas by kob
has nothing to do with food. Many other examples of territorial defence, including fish, birds or even
invertebrates, are related to competition for mates or safe lairs, rather than food. Territoriality will
emerge where there is a focused resource that provides enough for the individual or group, within a
boundary that is small enough to be defended without the expenditure of too much effort.
Many birds, particularly seabirds, though they nest in dense communities, are none the less territorial
in that they defend their nesting site to within the distance that they can reach while brooding. This is
necessary to prevent attacks on their own chicks or nesting material from neighbours. Commonly the
resulting superimposition of the short-range repulsion onto the long-range attraction characteristically
leads to the well-known roughly hexagonal spacing of nests. Interestingly, one gets a similar
hexagonal spacing resulting from the territorial behaviour of gardening limpets such as species of
Scutellastra . They vigorously defend their gardens of particular species of algae, that extend for
perhaps 1–2 cm around the periphery of their shells.
Territoriality is least likely with insectivorous birds, where the food supply is plentiful but
unpredictably distributed. Swifts rarely defend an area larger than the nest. Conversely, other
insectivorous birds that occupy more constrained territories, such as the ground-nesting Blacksmith
Lapwing may be very territorial, especially in the breeding season, where they not only threaten or
attack many kinds of intruders, but have stereotyped display behaviour to deter conspecifics sharing
neighbouring nesting spots.
Conversely, large solitary (or paired) carnivores, such as bears and the bigger raptors require an
extensive protected area to guarantee their food supply. This territoriality will only break down when
there is a glut of food, for example when Grizzly Bears are attracted to migrating salmon.
Animal sexual behaviour
Animal sexual behaviour takes many different forms, even within the same species. Researchers
have observed monogamy, promiscuity, sex between species, sexual arousal from objects or places,
sex apparently via duress or coercion, copulation with dead animals, homosexual, heterosexual and
bisexual sexual behaviour, and situational sexual behaviour and a range of other practices among
animals other than humans. Related studies have noted diversity in sexed bodies and gendered
behaviour, such as intersex and transgender animals.
The study of animal sexuality (and primate sexuality especially) is a rapidly developing field. It used
to be believed that only humans and a handful of other species performed sexual acts other than for
procreation, and that animals' sexuality was instinctive and a simple response to the "right"
stimulation (sight, scent). Current understanding is that many species that were formerly believed
monogamous have now been proven to be promiscuous or opportunistic in nature; a wide range of
species appear both to masturbate and to use objects as tools to help them do so; in many species
animals try to give and get sexual stimulation with others where procreation is not the aim; and
homosexual behaviour has now been observed among 1,500 species and in 500 of those it is well
documented.
Mating systems
In sociobiology and behavioural ecology, the term mating system is used to describe the ways in
which animal societies are structured in relation to sexual behaviour. The mating system specifies
which males mate with which females, and under what circumstances.
The following are some of the mating systems generally recognised in humans and other animals:
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• Monogamy: One male and one female have an exclusive mating relationship.
• Polygamy: A single individual concurrently carries a relationship/mates with one or more of
the opposite sex. Three types are recognized:
○ Polygyny (the most common polygamous mating system in vertebrates so far studied):
One male has an exclusive relationship with two or more females.
○ Polyandry: One female has an exclusive relationship with two or more males.
○ Polygynandry: Two or more individuals have an exclusive relationship with two or
more individuals from the opposite sex; the numbers of males and females need not be
equal, and in vertebrate species studied so far, there are usually fewer males.
• Promiscuity: Any male and female will mate within the social group.
Monogamy
Zoologists and biologists now have solid evidence that monogamous pairs of animals are not always
sexually exclusive. Many animals that form pairs to mate and raise offspring regularly engage in
sexual activities with extra-pair partners. This includes previous exemplars such as swans. Sometimes
these extra-pair sexual activities lead to offspring. Genetic tests frequently show that some of the
offspring raised by a monogamous pair come from the female mating with an extra-pair male partner.
These discoveries have led biologists to adopt new ways of talking about monogamy:
Social monogamy
Social monogamy refers to a male and female's social living arrangement (e.g., shared use of a
territory, behaviour indicative of a social pair, and/or proximity between a male and female) without
inferring any sexual interactions or reproductive patterns. In humans, social monogamy equals
monogamous marriage. Sexual monogamy is defined as an exclusive sexual relationship between a
female and a male based on observations of sexual interactions. Finally, the term genetic monogamy
is used when DNA analyses can confirm that a female-male pair reproduce exclusively with each
other. A combination of terms indicates examples where levels of relationships coincide, e.g.,
sociosexual and sociogenetic monogamy describe corresponding social and sexual, and social and
genetic monogamous relationships, respectively.
Whatever makes a pair of animals socially monogamous does not necessarily make them sexually or
genetically monogamous. Social monogamy, sexual monogamy, and genetic monogamy can occur in
different combinations.
Social monogamy is relatively rare in the animal kingdom. The actual incidence of social monogamy
varies greatly across different branches of the evolutionary tree. Over 90 percent of avian species are
socially monogamous. This stands in contrast to mammals. Only 3 percent of mammalian species are
socially monogamous, although up to 15 percent of primate species are socially monogamous. Social
monogamy has also been observed in reptiles, fish, and insects.
Sexual monogamy is also rare among animals. Many socially monogamous species engage in extra-
pair copulations, making them sexually non-monogamous. For example, while over 90% of birds are
socially monogamous, "on average, 30 percent or more of the baby birds in any nest [are] sired by
someone other than the resident male." Patricia Adair Gowaty has estimated that, out of 180 different
species of socially monogamous songbirds, only 10 percent are sexually monogamous.
The incidence of genetic monogamy, determined by DNA fingerprinting, varies widely across
species. For a few rare species, the incidence of genetic monogamy is 100 percent, with all offspring
genetically related to the socially monogamous pair. But genetic monogamy is strikingly low in other
species. Barash and Lipton note:
The highest known frequency of extra-pair copulations are found among the fairy-wrens, lovely
tropical creatures technically known as Malurus splendens and Malurus cyaneus. More than 65
percent of all fairy-wren chicks are fathered by males outside the supposed breeding group.
Such low levels of genetic monogamy have surprised biologists and zoologists, forcing them to
rethink the role of social monogamy in evolution. They can no longer assume social monogamy
determines how genes are distributed in a species. The lower the rates of genetic monogamy among
socially monogamous pairs, the less of a role social monogamy plays in determining how genes are
distributed among offspring. See also Evolution of Monogamy.
Polygamy
Polygamy is defined as a mating structure in which a single individual of one gender has exclusive
access to several individuals of the opposite gender. It takes two main forms - polygyny and
polyandry. As polygyny is the most common form of polygamy among vertebrates (including
humans, to some extent), it has been studied far more extensively than polyandry.
Polygyny
In some species, notably those with harem-like structures, only one of a few males in a group of
females will mate. Technically, polygyny in sociobiology and zoology is defined as a system in

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which a male has a relationships with more than one female, but the females are predominantly
bonded to a single male. Should the active male be driven out, killed, or otherwise removed from the
group, in a number of species the new male will ensure that breeding resources are not wasted on
another male's young. The new male may achieve this in many different ways, including:
• Competitive infanticide
in lions and some monkeys, the new male will kill the offspring of the previous alpha male to
cause their mothers to become receptive to his sexual advances since they are no longer
nursing.
• Harassment to miscarriage
amongst wild horses and baboons, the male will "systematically harass" pregnant females
until they miscarry.
• Pheromone based spontaneous abortion
in some rodents such as mice, a new male with a different scent will cause females who are
pregnant to spontaneously fail to implant recently fertilized eggs. This does not require
contact; it is mediated by scent alone. It is known as the Bruce-Parkes effect.
Promiscuity
Two examples of systems in primates are promiscuous mating chimpanzees and bonobos. These
species live in social groups consisting of several males and several females. Each female copulates
with many males, and vice versa. In bonobos, the amount of promiscuity is particularly striking
because bonobos use sex to alleviate social conflict as well as to reproduce.
Seasonal nature of animal sexuality
Many animal species have specific mating (or breeding) seasons (seasonal breeding). These are often
associated with changes to herd or group structure, and behavioural changes, including territorialism
amongst individuals. These may be annual (e.g. wolves), biannual (e.g. dogs) or more frequently (e.g.
horses). During these periods, females of most species are more mentally and physically receptive to
sexual advances, a period scientifically described as estrous but commonly described as being "in
season" or "in heat", but outside them animals still engage in sexual behaviours, and such acts as do
occur are not necessarily harmful.
Interpretation of animal sexuality
Many researchers have described homosexuality as something altogether different from sex. They
must realise that animals can have sex with who they will, when they will and without consideration
to a researcher's ethical principles.
When nine out of ten pairings occur between males, "[e]very male that sniffed a female was reported
as sex, while anal intercourse with orgasm between males was only [categorized as] 'revolving
around' dominance, competition or greetings.
Sex for pleasure
It is a common myth that animals do not (as a rule) have sex for pleasure, or alternatively that
humans, pigs (and perhaps dolphins and one or two species of primate) are the only species which do.
This is sometimes formulated "animals mate only for reproduction".
Of course, we have to make many seemingly artificial distinctions to arrive at our conclusion.
Animals other than humans have no awareness that their sexual activities are connected with
reproduction: They engage in sex because they're biologically driven to do so, and if the fulfillment of
their urges produces a physical sensation we might appropriately call 'pleasure,' it isn't the least bit
affected by the possibility (or impossibility) of producing offspring. We are also discounting cases in
which animals do engage in sex even though reproduction is an impossibility because we claim there
are other 'purposes' (of which the animals themselves are unaware) at play. (For example, the females
of some species of birds will invite males to mate with them even after they have laid their eggs, but
we ascribe a purpose to this behaviour: this is a biological "trick" to fool males into caring for
hatchlings they didn't father.) We also employ subjective terms such as 'willingly' and 'regularly' in
claiming that bonobos and dolphins are the only other animals who "willingly (and regularly) engage
in sex with each other" ... and even then it may be the case that these species have some other
'purpose' for doing so that we haven't yet discovered.
A 2006 Danish Animal Ethics Council report which examined current knowledge of animal sexuality
in the context of legal queries concerning sexual acts by humans, has the following comments,
primarily related to domestically common animals:
Even though the evolution-related purpose of mating can be said to be reproduction, it is not actually
the creating of offspring which originally causes them to mate. It is probable that they mate because
they are motivated for the actual copulation, and because this is connected with a positive experience.

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It is therefore reasonable to assume that there is some form of pleasure or satisfaction connected with
the act. This assumption is confirmed by the behaviour of males, who in the case of many species are
prepared to work to get access to female animals, especially if the female animal is in oestrus, and
males who for breeding purposes are used to having sperm collected become very eager, when the
equipment they associate with the collection is taken out.
There is nothing in female mammals’ anatomy or physiology, that contradicts that stimulation of the
sexual organs and mating is able to be a positive experience. For instance, the clitoris acts in the same
way as with women, and scientific studies have shown that the success of reproduction is improved
by stimulation of clitoris on (among other species) cows and mares in connection with insemination,
because it improves the transportation of the sperm due to contractions of the inner genitalia. This
probably also concerns female animals of other animal species, and contractions in the inner genitals
are seen e.g. also during orgasm for women. It is therefore reasonable to assume that sexual
intercourse may be linked with a positive experience for female animals.
Homosexual behaviour
Two male Mallards, Anas platyrhynchos. Mallards form male-female pairs only until the female lays
eggs, at which time the male leaves the female. Mallards have rates of male-male sexual activity that
are unusually high for birds, in some cases, as high as 19% of all pairs in a population.
The presence of same-sex sexual behaviour was not scientifically observed on a large scale until
recent times. Homosexual behaviour does occur in the animal kingdom outside humans, especially in
social species, particularly in marine birds and mammals, monkeys, and the great apes. Homosexual
behaviour has been observed among 1,500 species, and in 500 of those it is well documented.
Approximately eight percent of [male] rams exhibit sexual preferences [that is, even when
given a choice] for male partners (male-oriented rams) in contrast to most rams, which prefer
female partners (female-oriented rams). We identified a cell group within the medial preoptic
area/anterior hypothalamus of age-matched adult sheep that was significantly larger in adult
rams than in ewes...
• Male bighorn sheep are divisible into two kinds: the typical males among whom homosexual
behaviour, including intercourse, is common and "effeminate sheep", or "behavioural
transvestites", which are not known to engage in homosexual behaviour.
Cross species sex
While it is commonly believed that animal sexuality is instinctive and thus somewhat mechanistic,
research regularly records that many animals are sexual opportunists, partaking in sexual relations
with individuals of visibly distinct species. This is more visible in domesticated species and animals
in captivity, as domestication commonly selects for increased breeding rate (and so an accelerated
breeding cycle has commonly arisen in domesticated species over the centuries), and also because
these species are more easily observed by humans. Nevertheless, animals have been observed in the
wild to attempt sexual activity with other species or indeed inanimate objects. Attempts by wild
moose to obtain sex from domestic horses are apparently well known by wildlife specialists. In the
wild, where observation is harder, genetic studies have shown a "large number" of inter-species
hybrids, and other investigations describe productive and non-productive inter-species mating as a
"natural occurrence". Recent genetic evidence strongly suggesting this has occurred even within the
history of the human species, and that early humans often had sexual activity with other primate
species, is considered below.
Hybrid offspring can result from two organisms of distinct but closely related parent species,
although the resulting offspring is not always fertile. According to the definition of a species, if two
organisms cannot or will not mate and produce a fertile offspring, they are different species. The
mule, for example (a horse/donkey cross) is normally sterile, whilst the liger (lion/tiger cross) has
fertile females and sterile males Novosibirsk zoo director Rostislav Shilo says on the liger (born in its
zoo); "It’s just that the lion and the tiger live in neighboring caves in the Novosibirsk zoo, and got
used to each other. It’s practically impossible in the wild.
Due to the difficulties of observation, interspecies sex of this kind between two top-level predators,
occurring in the wild, was only conclusively documented with the finding of a grizzly-polar bear
hybrid in April 2006. Again, as with lions and tigers, the two species would normally not share
enough common territory to provide adequate opportunity for much cross-species sexual activity.
Prostitution
In some penguin species, the females, even when in a committed relationship, will exchange sexual
favours with strange males for the pebbles they need to build their nests. Prostitution was also
observed among chimpanzees, who trade food for sex.

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Sexual fetishes
Although not often reported, animals, or primates at the least, are able to sexualize inanimate objects
similar to the way human beings sexualize the objects of their sexual fetishes. Not only will an animal
that has a habitual object for masturbation sometimes appear to sexualize that object, primates have
generalized further to sexualize kinds of objects for which no instinctual or prior sexual connection
exists.
The sexualization of objects or locations is also well recognized in the breeding world. So for
example, stallions may often 'drop' (become sexually aroused) upon visiting a location where they
have been allowed to have sex before, or upon seeing a stimulus previously associated with sexual
activity such as an artificial vagina.
In this case however, the primary structure is Pavlovian conditioning, and the fetishistic association is
due to a conditioned response (or association) formed with a distinctive 'reward'. Human fetishism
can also be traced back to similar or near-identical conditioning: likewise based upon the Pavlovian
association between an erotic sensation or anticipation, and objects which become mentally
associated with that activity.
Sexual imagery viewing
A study by Platt, Khera and Deaner at Duke University North Carolina, showed that male monkeys
will give up privileges (in this case, juice, which is highly valued), to be allowed to see a female
monkey's hindquarters. Deaner and his team reported that monkeys would take a juice cut to look at
powerful males' faces or the perineum of a female, but to persuade the monkeys to stare at
subordinate males, the researchers had to bribe them with larger drinks. "Virtually all [male] monkeys
will give up juice to see female hindquarters ... they really value the images."
The researchers stress that in monkey society, such behaviours have great social utility and we should
therefore not simply reach the conclusion that "monkeys enjoy pornographic pictures". There is no
evidence at this point that viewable pictures or movies of sexual activity are valued for their sexual
enjoyment, although as noted above (Masturbation), there are reports that watching sex in real life
may have such an effect. The subject of animals and sexual imagery is not yet well researched.
Problems with encouraging pandas to mate in captivity have been very common. However, showing
young male pandas "panda pornography" is widely credited with a recent population boom among
pandas in zoos.
Coercive sex
Controversial interpretations and implications aside, sex in a forceful or apparently coercive context
has also been documented in a variety of species. A notable example is bottlenose dolphins, where at
times, a pod of bachelor males will 'corner' a female. Furthermore, in a zoo where it is common
practice to put newly captured dolphins in with dolphins who are established in their enclosures, other
species of dolphin are never put in together with bottlenoses because the bottlenose dolphins
frequently torment and rape them. The behaviour is also common in some arachnids (spiders),
notably those whose females eat the males during sex if not tricked with food and/or tied down with
threads, and in some herbivorous herd species or species where males and females are very different
in size, where the male dominates sexually by sheer force and size. Some species of birds appear to
combine sexual intercourse with apparent violent assault; these include ducks, geese, and white-
fronted bee-eaters. According to Emlen and Wrege (1986) forced copulations occur in this socially
nesting species, and females must avoid the unwelcome attention of males as they emerge from their
nest burrows or they are forced to the ground and mated with. Apparently, such attacks are made
preferentially on females who are laying and who may thus mother their offspring as a result.
In 2007, research suggested that in the Acilius genus of water beetles (also known as "diving
beetles"), an "evolutionary arms race" between the genders means that there is no courtship system
for these beetles. "It's a system of rape. But the females don't take things quietly. They evolve
counter-weapons." Cited mating behaviours include males suffocating females underwater till
exhausted, and allowing only occasional access to the surface to breathe for up to six hours (to
prevent them breeding with other males), and females which have a variety of body shapings (to
prevent males from gaining a grip). Foreplay is "limited to the female desperately trying to dislodge
the male by swimming frantically around."
Since the early 1990’s, for example, young male elephants in Pilanesberg National Park and the
Hluhluwe-Umfolozi Game Reserve in South Africa have been raping and killing rhinoceroses; this
abnormal behaviour, according to a 2001 study in the journal Pachyderm, has been reported in ‘‘a
number of reserves’’ in the region.
Sex between adults and juveniles
It has also been recorded that certain species of mole will impregnate newborns of their own species.
It is not clear if this is forceful or not. Similarly, the male stoat (Mustela erminea) will mate with
infant females of their species. This apparently is a natural part of their reproductive biology - there is
a delayed gestation period, so these females give birth the following year when they are fully grown.
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A male spotted hyena which attempted to mate with a female which succeeded in driving it off,
eventually turned to its ten-month-old cub, repeatedly mounting it and ejaculating on it. The cub
sometimes ignored this and sometimes struggled 'slightly as if in play'. The mother did not intervene.
Infants and children in Bonobo societies are often involved in sexual behaviour.
Sexual cannibalism
Sexual cannibalism, which has been documented in arachnids, insects and amphipods, is a
phenomenon in which a female organism kills and consumes the male before, during, or after
copulation. Although it does confer some known advantages to reproduction, whether or not the male
is complicit has not been scientifically determined.
Necrophilia
Necrophilia in animals is where a living animal engages in a sexual act with a dead animal. In one of
the most well-known examples, Kees Moeliker of the Rotterdam Natural History Museum,
Netherlands observed sexual activities outside his office between a live duck and a dead one. Two
male mallards which Moeliker believed were engaged in rape flight, a common motif in duck sexual
behaviour, collided with his window. "When one died the other one just went for it and didn't get any
negative feedback—well, didn't get any feedback," according to Moeliker, who described the event as
"homosexual necrophilia." The case was reported scientifically in Deinsea 8-2001, along with
photos., and earned Moeliker an Ig Nobel Prize in biology, awarded for research that cannot or should
not be reproduced.
Additionally, male cane toads have been documented (in Cane Toads: An Unnatural History)
engaging in copulation with dead toads and inanimate objects.
Notes on specific species
Bonobos
The Bonobo, which has a matriarchal society, is a fully bisexual species — both males and females
engage in sexual behaviour with the same and the opposite sex, with females being particularly noted
for engaging in sexual behaviour with each other and at up to 75% of sexual activity being
nonreproductive. Primatologist Frans de Waal believes that Bonobos use sexual activity to resolve
conflict between individuals. Sexual activity occurs between almost all ages and sexes of Bonobo
societies.
Birds
Some black swans of Australia form sexually active male-male mated pairs and steal nests, or form
temporary threesomes with females to obtain eggs, driving away the female after she lays the eggs.
More of their cygnets survive to adulthood than those of different-sex pairs possibly due to their
superior ability to defend large portions of land.
In early February 2004 the New York Times reported that a male pair of chinstrap penguins named
Roy and Silo in the Central Park Zoo in New York City were partnered and had successfully hatched
a female chick from an egg. Other penguins in New York have also been reported to be forming
same-sex pairs.
Zoos in Japan and Germany have also documented male penguin couples. The couples have been
shown to build nests together and use a stone to replace an egg in the nest. Researchers at Rikkyo
University in Tokyo, found twenty such pairs at sixteen major aquariums and zoos in Japan.
Bremerhaven Zoo in Germany attempted to break up the male couples by importing female penguins
from Sweden and separating the male couples; they were unsuccessful. The zoo director stated the
relationships were too strong between the couples.
Recently, a mated pair of swans in Boston were found to both be female. They too had attempted to
raise eggs together.
Studies have shown that ten to fifteen percent of female western gulls in some populations in the wild
prefer other females.
As many as 19% of Mallard pairs in a given population have been observed to consist of male-male
homosexuals.
Lizards
Whip-tailed lizard females have the ability to reproduce through parthenogenesis and as such males
are rare and sexual breeding non-standard. Females engage in sexual behaviour to stimulate
ovulation, with their behaviour following their hormonal cycles; during low levels of estrogen, these
(female) lizards engage in "masculine" sexual roles. Those animals with currently high estrogen
levels assume "feminine" sexual roles.
Lizards that perform the courtship ritual have greater fecundity than those kept in isolation due to an
increase in hormones triggered by the sexual behaviours. So, even though asexual whiptail lizards
populations lack males, sexual stimuli still increase reproductive success.
From an evolutionary standpoint these females are passing their full genetic code to all of their
offspring rather than the 50% of genes that would be passed in sexual reproduction. Certain species of
gecko also reproduce by parthenogenesis.
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Flatworm
Penis fencing is a mating behaviour engaged in by certain species of flatworm, such as
Pseudobiceros bedfordi. Species which engage in the practice are hermaphroditic, possessing both
eggs and sperm-producing testes.
The species "fence" using two-headed dagger-like penises which are pointed, and white in color. One
organism inseminates the other. The sperm is absorbed through pores in the skin, causing
fertilization.
Sheep
An October 2003, study by Dr. Charles E. Roselli et al. (Oregon Health & Science University) states
that homosexuality in male sheep (found in eight percent of rams) is associated with a region in the
rams' brains which the authors call the "ovine Sexually Dimorphic Nucleus" (oSDN) which is half the
size of the corresponding region in other male sheep.
However, some view this study to be flawed in that the determination of homosexuality within the
sheep, (sample population of twenty-seven for the study), was to have animals who were unable to
mount female ewes placed in a cage with two stanchioned males and two unstanchioned females (that
is, the males could not move or struggle while the females could). Given the aggressive nature of the
sheep copulation, the uneven treatment of males and females, many see this as simply evidence that
the sheep in question were unable to be aggressive enough to mount females. Some say that the
results were situational sexuality, unlike the bonds seen in human homosexuality.
The scientists found that, "The oSDN in rams that preferred females was significantly larger and
contained more neurons than in male-oriented rams and ewes. In addition, the oSDN of the female-
oriented rams expressed higher levels of aromatase, a substance that converts testosterone to
estradiol, an estrogen hormone believed to facilitate typical male sexual behaviours. Aromatase
expression was no different between male-oriented rams and ewes."
"The dense cluster of neurons that comprise the oSDN express cytochrome P450 aromatase.
Aromatase mRNA levels in the oSDN were significantly greater in female-oriented rams than in
ewes, whereas male-oriented rams exhibited intermediate levels of expression." These results suggest
that "...naturally occurring variations in sexual partner preferences may be related to differences in
brain anatomy and its capacity for estrogen synthesis." As noted previously, given the potential
unagressiveness of the male population in question, the differing aromatase levels may also have been
evidence of aggression levels, not sexuality. The results of this study have not been confirmed by
others.
Spotted Hyena
The female Spotted Hyena has a unique urinary-genital system, closely resembling the penis of the
male, called a pseudo-penis. The family structure is matriarchal and dominance relationships with
strong sexual elements are routinely observed between related females.
They are notable for using visible sexual arousal as a sign of submission and not dominance, in males
as well as females (females have a sizable erectile clitoris), to the extent that biologist Robert
Sapolsky speculates that in order to facilitate this, their sympathetic and parasympathetic nervous
systems may be partially reversed in respect to their reproductive organs.
Bottlenose Dolphins
Bottlenose Dolphin males have been observed working in pairs to follow and/or restrict the
movement of a female for weeks at a time, waiting for her to become sexually receptive. The same
pairs have also been observed engaging in intense sexual play with each other.
Janet Mann, a professor of biology and psychology at Georgetown University, argues that the
common same-sex behaviour among male dolphin calves is about bond formation and benefits the
species evolutionarily. They cite studies that have shown the dolphins later in life are bisexual and the
male bonds forged from homosexuality work for protection as well as locating females with which to
reproduce.
In 1991 an English man was prosecuted for allegedly having sexual contact with a dolphin. The man
was found not guilty after it was revealed at trial that the dolphin was known to tow bathers through
the water by hooking its large penis around them.
Seahorses
Seahorses, long upheld as monogamous and mating for life, are identified as "promiscuous, flighty,
and more than a little bit gay" according to research published in 2007.
Scientists at 15 aquariums studied 90 seahorses of 3 species. Of 3168 sexual encounters, 37% were
same sex acts. Flirting was common (up to 25 potential partners a day of both genders); only one
species (the British Spiny Seahorse) included faithful representatives, and for these 5 of 17 were
faithful, 12 were not. Bisexuality was widespread and considered "both a great surprise and a shock",
with big bellied seahorses of both genders not showing partner preference. 1986 contacts were male-
female, 836 were female-female and 346 were male-male.

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Lions
Male lions often lead their social groups jointly with one or more of their brothers. To ensure loyalty,
the male co-leaders will "strengthen the bonds" by often having sex with each other.
Horses
Anecdotal evidence suggest that some horses have environment or appearance preferences when
selecting mates. There is also anecdotal evidence of limited bisexual behaviour in some stallions,
although there is (as of 2008) no conclusive scientific confirmation. The anecdotal evidence claims
this is most likely to occur in a single isolated group, with no access to mares.
Humboldt penguins
In 2009 at a zoo in Bremerhaven, Germany, two male adult humboldt penguins adopted an egg that
had been abandoned by its biological parents. After the egg hatched, the two penguins raised,
protected, cared for, and fed the chick in the same manner that heterosexual penguins raise their own
biological offspring.
Other evidence of interspecies sexual activity
Looking back in history, current research into human evolution tends to confirm that in some cases,
interspecies sexual activity may have been responsible for the evolution of entire new species.
Analysis of human and animal genes in 2006 provides strong evidence that after humans had
diverged from other apes, interspecies mating nonetheless occurred regularly enough to change
certain genes in the new gene pool:
A new comparison of the human and chimp genomes suggests that after the two lineages separated,
they may have begun interbreeding. [...] A principal finding is that the X chromosomes of humans
and chimps appear to have diverged about 1.2 million years more recently than the other
chromosomes.
The research suggests that:
There were in fact two splits between the human and chimp lineages, with the first being followed by
interbreeding between the two populations and then a second split. The suggestion of a hybridization
has startled paleoanthropologists, who nonetheless are "treating the new genetic data seriously."
The Washington Post comments, "If this theory proves correct, it will mean modern people are
descended from something akin to chimp-human hybrids."
A 1932 exploitation film, The Sex Life of a Gorilla, was based on supposed reports of primitive cults
in Africa routinely having sexual relations with gorillas.
Instinct
Instinct are the inherent inclination of a living organism toward a particular behavior. The fixed
action patterns are unlearned and inherited. The stimuli can be variable due to imprinting in a
sensitive period or also genetically fixed. Examples of instinctual fixed action patterns can be
observed in the behavior of animals, which perform various activities (sometimes complex) that are
not based upon prior experience, such as reproduction, and feeding among insects. Sea turtles,
hatched on a beach, automatically move toward the ocean, and honeybees communicate by dance the
direction of a food source, all without formal instruction. Other examples include animal fighting,
animal courtship behavior, internal escape functions, and building of nests. Another term for the same
concept is innate behavior.
Instinctual actions - in contrast to actions based on learning which are served by memory and which
provide individually stored successful reactions built upon experience - have no learning curve, they
are hard-wired and ready to use without learning. Some instinctual behaviors depend on maturational
processes to appear.
Biological predispositions are innate and biologically-vectored behaviors that can be easily learned.
For example in one hour, a baby colt can learn to stand, walk, glide, skip, hop and run. A biological
predisposition may also mean that a person, because of his/her genetic makeup, is more prone to
certain conditions or disease. Learning is required to fine tune the neurological wiring reflex like
behavior. True reflexes can be distinguished from instincts by their seat in the nervous system;
reflexes are controlled by spinal or other peripheral ganglia, but instincts are the province of the brain.
In a situation when two instincts contradict each other, an animal may resort to a displacement
activity.
One theory on instinct is that the inherent inclination is stored in the DNA and therefore passed from
parent to offspring.
Evolution
Instinctive behavior can be demonstrated across much of the broad spectrum of animal life.
According to Darwin's theory of evolution by natural selection, a favorable trait, such as an instinct,
will be selected for through competition and improved survival rate of life forms possessing the
instinct. Thus, for evolutionary biology, instincts can be explained in terms of behaviors that increase
the reproductive success of the individual or its close relatives.
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A good example of an immediate instinct for certain types of bird is imprinting. This is the behaviour
that causes geese to follow around the first moving object that they encounter, as it tends to be their
mother. Much work was done on this concept by the psychologist Konrad Lorenz. Another important
aspect of imprinting is sexual selection. Young birds tend to prefer the features of the parent of the
opposite sex when it come to the preferred mate. An inhibitory effect is the Westermarck Effect, the
tendency not to be sexually attracted to siblings, because of the close contact in a sensitive period.
The idea that no learning is required for instinctive behavior does not always apply when it concerns
the behaviors themselves. But the key stimuli and the specific outcome may vary somewhat due to
different inputs. The output may vary in the sense that a finch will sing naturally (instinctively), but it
will sing a song similar to the songs it has picked up in a sensitive period which explains the different
regional accents in the finch songs.
The Baldwin Effect
The Baldwin effect functions in two steps. First, phenotypic plasticity allows an individual to adjust
to a partially successful mutation, which might otherwise be utterly useless to the individual. If this
mutation adds to inclusive fitness, it will succeed and proliferate in the population. Phenotypic
plasticity is typically very costly for an individual; learning requires time and energy, and on occasion
involves dangerous mistakes. Therefore there is a second step: provided enough time, evolution may
find an inexorable mechanism to replace the plastic mechanism. Thus a behavior that was once
learned (the first step) may in time become instinctive (the second step). At first glance, this looks
identical to Lamarckian evolution, but there is no direct alteration of the genotype, based on the
experience of the phenotype.
The term "instincts" has had a long and varied use in psychology. In the 1870s, Wilhelm Wundt
established the first psychology laboratory. At that time, psychology was primarily a branch of
philosophy, but behavior became increasingly examined within the framework of the scientific
method. While use of the scientific method led to increasingly rigorous definition of terms, by the
close of the 19th century most repeated behavior was considered instinctual. In a survey of the
literature at that time, one researcher chronicled 4000 human instincts, meaning someone applied the
label to any behavior that was repetitive. As research became more rigorous and terms better defined,
instinct as an explanation for human behavior became less common. In a conference in 1960, chaired
by Frank Beach, a pioneer in comparative psychology and attended by luminaries in the field, the
term was restricted in its application. During the 60's and 70's, textbooks still contained some
discussion of instincts in reference to human behavior. By the year 2000, a survey of the 12 best
selling textbooks in Introductory Psychology revealed only one reference to instincts, and that was in
regard to Sigmund Freud's referral to the "id" instincts.
Any repeated behavior can be called "instinctual," as can any behavior for which there is a strong
innate component. However, to distinguish behavior beyond the control of the organism from
behavior that has a repetitive component we can turn to the book "Instinct" (1961) stemming from the
1960 conference. A number of criteria were established which distinguishes instinctual from other
kinds of behavior. To be considered instinctual a behavior must a) be automatic, b) be irresistible, c)
occur at some point in development, d) be triggered by some event in the environment, e) occur in
every member of the species, f) be unmodifiable, and g) govern behavior for which the organism
needs no training (although the organism may profit from experience and to that degree the behavior
is modifiable). The absence of one or more of these criteria indicates that the behavior is not fully
instinctual. Instincts do exist in insects and animals as can be seen in behaviors that cannot be
changed by learning. Psychologists do recognize that humans do have biological predispositions or
behaviors that are easy to learn due to biological wiring, for example walking and talking.
If these criteria are used in a rigorous scientific manner, application of the term "instinct" cannot be
used in reference to human behavior. When terms, such as mothering, territoriality, eating, mating,
and so on, are used to denote human behavior they are seen to not meet the criteria listed above. In
comparison to animal behavior such as hibernation, migration, nest building, mating and so on that
are clearly instinctual, no human behavior meets the necessary criteria. And even in regard to
animals, in many cases if the correct learning is stopped from occurring these instinctual behaviors
disappear, suggesting that they are potent, but limited, biological predispostions. In the final analysis,
under this definition, there are no human instincts.
In humans
Some sociobiologists and ethologists have attempted to comprehend human and animal social
behavior in terms of instincts. Psychoanalysts have stated that instinct refers to human motivational
forces (such as sex and aggression), sometimes represented as life instinct and death instinct. This use
of the term motivational forces has mainly been replaced by the term instinctual drives.
Instincts in humans can also be seen in what are called instinctive reflexes. Reflexes, such as the
Babinski Reflex (fanning of the toes when the foot is stroked), are seen in babies and are indicative of
stages of development. These reflexes can truly be considered instinctive because they are generally
free of environmental influences or conditioning.
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Additional human traits that have been looked at as instincts are: sleeping, disgust, face perception,
language acquisitions, "fight or flight" and "subjugate or be subjugated". Some experiments in human
and primate societies have also come to the conclusion that a sense of fairness could be considered
instinctual, with humans and apes willing to harm their own interests in protesting unfair treatment of
self or others.
Many scientists consider that it is instinctual in children to put everything in their mouths, because
this is how they tell their immune system about the environment and the surroundings, what the
immune system should adapt to.
Other sociologists argue that humans have no instincts, defining them as a "complex pattern of
behavior present in every specimen of a particular species, that is innate, and that cannot be
overridden." Said sociologists argue that drives such as sex and hunger cannot be considered
instincts, as they can be overridden. This definitory argument is present in many introductory
sociology and biology textbooks, but is still hotly debated.
Psychologist Abraham Maslow argued that humans no longer have instincts because we have the
ability to override them in certain situations. He felt that what is called instinct is often imprecisely
defined, and really amounts to strong drives. For Maslow, an instinct is something which cannot be
overridden, and therefore while it may have applied to humans in the past it no longer does.
In the book An Instinct for Dragons anthropologist David E. Jones suggests a hypothesis that humans
just like monkeys have inherited instinctive reactions to snakes, large cats and birds of prey. Folklore
dragons have features that are combinations of these three, which would explain why dragons with
similar features occur in stories from independent cultures on all continents. Other authors have
suggested that especially under the influence of drugs or in dreams, this instinct may give raise to
fantasies about dragons, snakes, spiders, which makes these symbols popular in drug culture. The
traditional mainstream explanation to the folklore dragons does however not rely on human instinct,
but on the assumption that fossil remains of dinosaurus gave raise to similar speculations all over the
world.
Memory
Stored information plays a critical role in the lives of many animals. Knowing a nest location,
landmarks for navigation in a home range, where food and water have been found the past, and how
previous social interactions with another animal have turned out, all are critical pieces of information
in shaping future behavior.
One of the most interesting aspects of the study of memory is that it has essentially the same
functional structure and underlying biochemistry across the animal kingdom. This suggests that
memory or its evolutionary predecessor evolved early in the history of the animals. Studies of
learning and memory in a sea slug, Aplysia, have resulted in a model of learning and memory that
seems to be broadly applicable to all animals. Eric Kandel of Columbia University was awarded the
Nobel Prize in 2000 for his work on learning and memory and for developing Aplysia as a model
system.
Memory starts as a biochemical response in the brain following sensory input. The first change comes
at the level of the synapse--junctions between nerve cells. Acetylcholine, a well known
neurotransmitter, plays a critical synaptic role in the initial formation of memory. We can think of
short-term memory as resulting from a transient change in neurostransmitter levels at synapses.
Treatments for memory-deficit problems, like Alzheimer's, in humans often involve enhancing the
retention of acetylcholine in brain synapses. Aricept (donepezil hydrochloride), a drug marketed to
prevent memory problems, acts by inhibiting the action of the enzyme in the brain that breaks down
acetylcholine (acetylcholinesterase). People with moderate levels of dementia can show marked
improvement with this sort of treatment.
At a behavioral level, what happens next depends on the animal's experience after the initial learning
phase. If the animal moves on and doesn't have immediate reinforcement of the learned event, likely
that memory quickly fades and is irrecoverable. Short-term memory is highly subject to amnesia
effects. A traumatic event can obliterate short-term memory, as can a variety experimental treatments,
such as carbon dioxide or nitrogen narcosis or chilling. This makes sense, as retention relies on
neurotransmitter levels, which can be quite sensitive to an animal's general internal physiological
state.
Evolutionarily, the transient nature of short-term memory is a huge advantage in allowing the animal
to sort learning that only has immediate importance from learning that will be useful in the days or
weeks ahead. We can think of the rapid turnover of information in short-term memory as a way of
continuously clearing the mental "chalkboard" of potentially extraneous information.
Some information, though, has critical future importance and needs to be moved to long-term
memory. Oftentimes reinforcement--re-experiencing the item to be learned--plays a critical role in
the formation of long-term memory. Another key behavioral aspect of the formation of long-term
memory is the formation of associations. Often associations are sensory; for example, humans talk
about odors which are evocative of complex sets of memories from the past. This combination of
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reinforcement and association can be thought of as helping to build a network of neurons which hold
this key information. Association is probably a key element in accessing long-term memory. A single
memory which is not linked to other remembered items is quite difficult to access. Forming long-term
memories requires the synthesis of proteins in neurons. If protein synthesis is inhibited or disrupted,
long-term memories will not be formed.
Between short-term and long-term memory lies middle-term memory, or the consolidation phase.
This is a time of transition between the neurotransmitter/synaptic basis for short-term memory and
the protein based long-term memory. At least in the honey bee, short-term memory and middle-term
memory are independent--both are formed during learning, but only middle-term memory is the
gateway to forming long term memories.
While treatments, like the Aricept mentioned above, can delay the onset of serious memory deficit
problems in humans, death of brain cells is also a key issue in human dementia. Once the cells have
died, treatment at the neurotransmitter level is futile. Treatments currently under development are
aimed at preventing cell death. There is outstanding promise, with a combination of a treatment that
manipulates acetylcholine levels and one that prevents cell death, for managing human brains to
prevent dementia.
Social Behavior
Behavior - the sum of all the motor responses of an organism to all the external and internal stimuli
acting upon it.
Social behavior of animals is the sum of inter-individual relationships among the members of that
population.

Mechanisms
Behaviors involved in interactions among species are agonistic behaviors, predator-prey behaviors
and reproductive behaviors.
These behaviors result in individual distance the "kick-kiss" distance.
Spacing patterns and social groupings
Spacing pattern

Social grouping Solitary Home Range Territorial

Solitary yes yes yes

Male-female pair yes yes yes

Small group yes yes yes

School/School "aggregation" yes yes no


Fish/groups may be randomly distrubuted or clumped as in home ranges, territories, and schools.
Home range; the area through which an animal or group regularly travels during daily activities, but
there may be a core area. The shift from random distributions to home ranges of solitary fishes may
relate to differences in resource richness.
Territory; an area occupied almost exclusively by one or more animals, by active repulsion of
potential intruders through defense or advertisement.
Advantages of a territory;
○ knowledge of territory, including refuges, good breeding sites etc.
○ access to mates.
○ access to food - within an economically defensible area.
Schools: groups of fishes range from aggregations to polarized schools. A polarized school has been
defined as relatively long-term association of three or more fish moving in concert at fixed distances.
Long-term associations, lacking rigid organization may be called non-polarized schools or shoals.
In schooling associations, there is no apparent behavioral differentiation among members, there are
no leaders, no persistent ties among near-neighbors and no dominance relationships. There is no overt
aggression, but inter-neighbor distances (individual distance) are conservative in polarized schools.
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Schooling behavior is very common among fish; Cyprinidae, Salmonidae, Clupeidae, Atherinidae,
Scombridae, Engraulidae. Most fish school as young but only about 20-25% as adults. Numbers in
school vary from a few individuals to millions in herring, menhaden. Atlantic herring schools
sometimes encountered in winter occupying 279 to 4580 m3 with 0.5 to 1 fish.m-3.
Behavioral patterns.
○ Short radius behavior.
○ Loose cruising association.
○ Predator encircling.
○ Flash expansion.
○ Mills.
Advantages of Schooling.
○ reduced risk of predation.
○ increased efficiency of finding food.
○ increased reproductive success.
○ increased swimming efficiency.
Factors Affecting School Structure
○ Threat.
○ time.
○ environment; presence of food, current etc.
○ life history stage.
○ Reproduction.
SOCIAL INTERACTION
Social interactions of various types are more important in determining degree of sociality than are
most of the above characteristics. The only interactions between animals that are seldom considered
social are behaviours in which animals take something needed by others. The question arises,
however, as to how to classify the parasitic relationships of a fetus in the mother and the male
anglerfish (Photocorynus spiniceps) on his mate, or the fact that killing by predators may help
animals to avoid overgrazing their habitats. One could speak of communication of feint and chase in
the interaction between moose and wolves. When a bird chases another off its territory, it uses
communication and interacts with the other bird very strongly.
Humans often consider chimpanzees or bees as more social than desert locusts, because the locusts
have rather simple interactions. Male hummingbirds and birds of paradise, however, have their
elaborate plumages and social displays because the females get together with them so seldom that
they might not recognize a suitable mate without the displays. It seems generally true that elaborate
rituals evolve where social bonds are most fleeting or likely to be disrupted. To determine sociality,
one must look at the total spectrum of social interactions as well as at their diversity and productivity,
rather than just at a single feature.
Altruistic social behaviour is often found among animals. An altruistic animal is one that expends
some of its energy helping another without direct benefit to itself, be it a mother bear protecting her
cubs against their hungry father or a bird giving an alarm call that warns its neighbours of a hawk. An
alarm call may help the bird itself, of course, by startling the predator or warning it that this alert bird
will be hard to catch.
Psychologists have found that a rat or monkey will slow its rate of pressing a lever for food if that
lever also gives an electric shock to a nearby rat or monkey. Rats will take turns sitting on a platform
so that others can feed without being interrupted by electric shock. Rats or pigeons can be trained to
cooperate in getting food.
Since the altruistic animal always loses something by its behaviour, the question arises why altruism
exists. One answer is that, as the evolutionist Charles Darwin suggested, when an animal protects its
offspring, it helps its kind to survive the process of natural selection. When porpoises help an injured
relative to the surface where it can breathe, they seem to be following a pattern of behaviour that can
be accounted for by evolution. Their altruism clearly helps the group and therefore becomes part of
the genetic endowment. Altruism, significantly enough, is usually limited to an animal's relatives.
Most social animals, such as penguins, feed only their own young. When the individual animal loses
more than it or its relatives gain, as when female seals nurse young not their own, the question arises
whether this serves the survival of the larger group or the species. Under some conditions, the
survival of the group may be more important even than survival of the individual, as when the
honeybee dies defending the hive. The worker honeybee, which is not able to reproduce, is in the
biological sense not an individual so much as an extra limb of a collective animal.
Reciprocal altruism, in which a benefit is later returned to the benefactor, need not be between related
animals and may not even seem altruistic. Alarm calls of birds often alert entirely unrelated kinds of
birds, which later may return the favour. An act that seems selfish in the short run is sometimes
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altruistic in the long run, or vice versa, in the case of maladaptation. Wasps, ants and termites that
cannibalize or dominate nestmates at times of food shortage may better keep the colony from
starving. Individual ants and bees are often lazy, spending most of their time resting or wandering
aimlessly, but these unemployed individuals form an easily mobilized reserve in times of danger.
Individual and group recognition are often important aspects of social structure. Ovenbirds (Seiurus
aurocapillus) in North America recognize neighbouring males by their songs and react aggressively
mainly to songs of strange males. Animals that have long parental bonds often show individual
recognition. Herring gulls (Larus argentatus), for example, recognize chicks or mates by slight
differences in voice or appearance. The larger or more ephemeral the society, the less there can be
individual recognition between distant individuals and the more important becomes recognition by
group characters, such as the "nest odours" of social insects. Ants, bees, and termites often attack
strangers, or even members of their own colony that have been experimentally removed for a few
days or washed. Many kinds of parasitic insects (beetles, flies, butterfly caterpillars), however,
provide food or scents that gain them entry to a nest, then prey on larvae there.
Other internal characteristics of societies are age structure, birth rates, and death rates. A young wasp
or termite colony has few old animals, a mature colony has more, and a declining colony or one that
is producing reproductive forms has few young. The old colony has a lower percentage of foraging
workers than does the young colony, and has a lower birth rate and higher death rate as a
consequence; but only the old colony produces reproductive forms.
Societies also perform movements, such as nomadism and migration (see above Migratory
behaviour). Army ants wander nomadically after prey. Wildebeest and locusts of Africa emigrate to
green areas of local rains; flocking or solitary birds migrate back and forth to escape winter or
drought; anadromous fishes, such as salmon, move to the sea for food and to rivers to spawn;
catadromous ones, such as eels, do the reverse.
Migrants are often placed at a disadvantage compared to residents, for the latter can take the regular
food supplies and leave only ephemeral sources for migrants. Migrants that follow army ants for food
in Central America are subordinate to residents and succeed only when residents are absent. The
migrant can turn its world from a coarse-grained one to a fine-grained or dependable one by
migrating from one patch to the next, and by force of numbers migrants sometimes displace residents.
Societies can make use of seasonal environmental changes by migrating locally, such as sowbugs
clustering to estivate in hot weather or ladybugs hibernating in masses. Other societies show food
storage; e.g., harvester ants (Messor) and wood or pack rats (Neotoma). Honey ants (Myrmecocystus)
have a "replete" caste that bloat their abdomens with stored honey and hang from the roofs of
underground chambers until tapped by other workers.
Societies show population fluctuations, from extinction to explosion. Mass emigrations in some, such
as lemmings and squirrels, occur mainly after population explosions or periodic extirpations of food
supplies. Some populations, such as many protozoans, worms, and insects, normally undergo violent
fluctuations, but are resistant to extinction. These are animals in which the adults emigrate or both
adults and young emigrate. Animals in which the adults normally occupy a fine-grained habitat and
only the young move, such as most higher animals, normally have relatively stable populations, but
are easily killed off by a new predator or temporally unpredictable events.
A major external characteristic of the more complex societies is that they construct things, or modify
their environments. The elaborate air-conditioned castles of some termites, the path systems of feral
house cats, and the patterns of singing at dawn among birds of ephemeral or coarse-grain habitats, all
are "structures" created by animals.
Cooperation and competition are major aspects of animal social behaviour. Social facilitation, as
when yawning spreads through a pride of lions or chickens eat more rapidly together, shows that
cooperative competition can be social. Social animals, which live close together, often interfere and
fight more with each other, especially in early stages, than do solitary ones. A fair percentage of the
communication between social animals involves "agonistic" or threat-submission behaviour. If this
behaviour results in a more adaptive dispersion of the animals, it has been altruistic. Grouped goldfish
and other animals survive heat, cold, metallic ions, and other "pollution" better than do isolated
animals; but if too many animals aggregate, they pollute each other. Social groups therefore have an
optimum size and density, based upon an equilibrium between advantages and disadvantages.
Linking the internal and external characteristics of social systems are flows of energy and materials.
Social systems use energy in building structures, or information-rich systems. Physically, one can
measure the success of a social system by how efficiently and extensively it uses energy and materials
and converts them to physical, biological, or cultural structures. Success in the short run can lead to
disaster in the long run, as when elephants or humans destroy an African forest and then must starve
or emigrate. Energy and material flows involve an interspecific web, the ecosystem, and must be
measured over the long run and in general as well as locally or in the short run.
Social behaviour, therefore, must include interactions between different kinds of animals. A flock of
sandpipers in flight is not less social if it includes two species rather than one. If species cooperate, a
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flock of two species can even be more social than a flock of one species. At African waterholes,
baboons keep the lookout while associated antelope are good at scenting predators. Symbiosis is
social, even though the late biologist Traian Savesculu of Romania was half right when he joked
"Symbiosis is like marriage--a mutual exploitation." Like all social organizations, it is both
cooperative and competitive.
Social behaviour may thus be defined as "more or less diverse and constructive interactions among
two or more animals." Social behaviour is usually constructive, productive, and adaptive; but it
sometimes persists for a time after the evolutionary basis for it is gone. For instance, many kinds of
hawks in the eastern United States have almost disappeared, but the flocks of small birds that were
formerly their prey still form as vigilante groups.
Which Animals are Eusocial?
Most eusocial animals are found in the phylum Arthropoda, but a few are found in the phylum
Chordata (Table 1).
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The order Hymenoptera is the largest and most well-known animal group with eusocial species. Most
hymenoptera are not eusocial, but this social system has arisen multiple times within the group. It is
found in some bees (Apidae) and wasps — with separate origins of eusociality in Sphecidae and
Vespidae — and all ants . Hymenoptera have a haplodiploid sex determination system (whereby
females arise from fertilized diploid eggs and males arise from unfertilized haploid eggs), which may
contribute to kin selection, favoring altruistic behavior in this group.
Termites can be considered to be highly evolved social cockroaches which live in their food. They
depend on a complex mutualism with cellulose digesting protozoans and bacteria, and young
individuals acquire these symbionts via anal trophallaxis. Within a colony, there is caste
differentiation: there is a queen and king, the sole reproducing individuals, and there are soldiers and
workers, which are different stages of pluripotent larvae of both sexes. Termites are diploid and
perform complex social behaviors including nest construction and territorial defense .
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The more recently discovered eusocial organisms include a few species of shrimp, aphids, and thrips.
There are at least two separate origins of eusociality within the Synalpheus shrimps . These marine
shrimp live in groups of several hundred closely related diploid individuals as internal parasites on
tropical sponges. The host sponges have a heterogeneous distribution, and this may have contributed
to the evolution of eusociality within this group as dispersal to establish new colonies is riskier than
remaining within the natal nest.
Thrips are small haplodiploid insects in the order Thysanoptera. Of about 5,000 species, about 300
form nests in plants called galls, feeding on plant tissue within. Of these, there are six species which
can be classified as eusocial: they have morphologically different soldier castes which defend the
galls from kleptoparasites.
Social aphids, like thrips, live in galls or hollowed stems of plants, feeding from the plant tissue
(Stern 1994, Aoki & Imai 2005). These tiny hemipterans may have complex life cycles — they are
diploid, but can reproduce parthenogenetically — and there have been several species described that
have robust soldier morphs .
There are at least two species of vertebrates that could be considered eusocial, the naked mole rat and
the Damaraland mole rat. Both species are diploid, highly inbred and live in harsh deserts with patchy
food resources. Most individuals help to raise siblings or close relatives that are born to a single
reproductive female (the queen). Outbreeding results from the generation of dispersive morphs —
individuals that have inbreeding avoidance and large fat stores — from the natal nest to start a new
colony.
Advantages to Living in Groups
Eusocial organisms live in groups, and are subject to both the benefits and costs of group living.
Living in a group may confer benefits on individuals in several ways. First, groups may form as
defense against predation, forming a "selfish herd" in which each individual has a lower chance of
being killed. Second, larger groups in many species gain advantages against competitors, such as in
colonies of the ant Azteca trigona.
Many eusocial organisms have strategic advantage when acquiring food in groups. Where there are
limited nest sites or resources are patchy, the benefits to staying within the natal group include
reducing dispersal risk and the possibility of inheriting the natal nest. This is a contributing factor in
naked mole rat societies and in insect societies with totipotent workers such as primitive ants and
wasps.
There can be costs to living in social groups, and these must ultimately be balanced by fitness
advantages. Disadvantages include increased competition for resources between individuals,
increased transmittance of parasites and diseases within groups, and easy detection of the group by
predators and parasites.
How did Eusociality Evolve?
Giving up one's reproductive potential is contrary to the basic premise of natural selection (to survive
and reproduce). Even Darwin (1859) commented on the challenge of understanding eusociality as
"one special difficulty, which at first appeared to me insuperable, and actually fatal to the whole
theory. I allude to the neuters or sterile females in insect communities: for these neuters often differ
widely in instinct and in structure from both the males and fertile females, and yet, from being sterile,
they cannot propagate their kind." Darwin goes on to argue that "This difficulty, though appearing
insuperable, is lessened, or, as I believe, disappears, when it is remembered that selection may be
applied to the family, as well as to the individual. . ." There have been several hypotheses proposed
for the evolution of worker-like behavior. It is important to note that they are not mutually exclusive
— each may play a different role in the evolution of eusociality in different groups. Evolutionary
biologists trace the origins of eusociality through a pathway that starts with solitary organisms
acquiring benefits to group behavior, eventually leading to a "point of no return" wherein certain
individuals no longer have the physical ability to reproduce and only gain evolutionary fitness
indirectly. In addition, it is important to note that the selective forces at work during the inception of
eusocial behavior may be different from those that maintain advanced eusocial colonies. What
follows is a brief description of major contributing factors during the inception of eusocial behavior.
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Figure 4: Diagram illustrating factors contributing to the evolution of primitive eusociality and that
advanced eusociality is a social state that is not reversible (past the point of "no return," individual
workers have lost the capacity to reproduce independently)
Kin Selection
There are two routes by which a gene can promote copies of itself in future generations: directly,
through producing offspring, and indirectly, through the reproduction of close relatives. The sum of
direct and indirect reproductive gains is known as inclusive fitness. As a result, it is possible for
selection of eusociality over solitary behavior if indirect fitness levels exceed direct fitness.
An altruistic act is one which benefits a recipient at a cost to the performer of the act. Hamilton's rule
(Hamilton 1964) states that altruism is favored if
r > C/B
where B is the benefit to recipient of altruistic act in terms of lifetime reproductive success and C is
the cost (decrease in lifetime reproductive success). The coefficient of relatedness, r, ranges from 0 to
1 and is the proportion of alleles shared between two individuals identical by descent. Eusociality
depends on high levels of altruism within groups as individuals increase the fitness of others at a cost
to themselves (e.g., by helping care for brood, defending nests, sharing food resources, not investing
in their own offspring).
There are two well-known mechanisms by which r can be elevated: haplodiploid sex determination
and inbreeding. While is it not required for eusociality to evolve (naked mole rats and termites are
diplodiploid), haplodiploidy may help to explain why eusociality has arisen multiple times within the
Hymenoptera. In haplodiploid organisms, the relatedness between full sibling sisters (r = 0.75) is
greater than between a mother and her offspring (r = 0.5), thus weighting Hamilton's rule in favor of
raising sisters rather than offspring.
Inbreeding, or mating between close relatives, results in offspring which share a larger proportion of
alleles, thus increasing r. This is common in organisms which don't disperse very long distances from
the natal nest or are prone to mating with siblings (e.g., termites and wild naked mole rats;
Delayed Benefits
An intermediate step toward eusociality may have "hopeful reproductives" — that, workers which
have the option to stay and help or to go out start their own nest. The decision can depend on
hierarchical position within the group, territory, or food resources and other environmental
conditions. Florida scrub jay offspring are known to stay at the natal nest, helping to raise siblings
(thus increasing their inclusive fitness) until there is an opportunity to replace their parents.
Primitively eusocial wasp colonies, such as Polistes, are commonly inherited by dominant workers on
the death of a queen.
Multi-level Selection
Natural selection can occur at the level of the individual, family (related individuals, known as kin),
or group (non-related individuals). This is known as multi-level selection. For eusocial organisms, the
traits (phenotype) of the colony interact with the environment to determine colony-level fitness and
can be described with models of multi-level or trait-group selection. Whether models of multi-level
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selection or inclusive fitness models are the best approach to explore the origin and maintenance of
eusociality remains a strong point of debate especially since there is limited empirical evidence for
inclusive fitness in groups .
Ecological and Life History Contributions
Nesting behavior has been described as a possible prerequisite for the development of eusociality, in
large part since it creates situations conducive to cooperative brood care . Where nest founding is
dangerous or there are limited territories or spaces, "fortress defenders" can cooperate to defend this
valuable resource. In the case of termites, thrips, shrimp, and aphids, the protected nest is also the
location of food in a patchy environment. Parental care can also be an important life history
component. One path to eusociality in Hymenoptera is thought to start with solitary females engaging
in simultaneous progressive provisioning — rearing multiple larvae of different ages at the same
time. Transitioning to eusocial behavior would then incorporate remaining offspring and provisioning
siblings, followed by offspring withholding their own reproduction.
Communication Among Animals
Communication is the transfer of information. Animals often communicate among their own species.
Sometimes they even try to communicate with other species. Most communication is done through
body language. Sounds and odors are also means to communicate. Humans can learn from how
animals communicate among each other, since the same methods apply in a subtle manner among
humans.
Questions you may have include:
• How do animals use body language?
• How do animals communicate with sounds and odors?
• What can humans learn from animals ?
• Communication Among Animal Species
• 1. Scent
• Smell is probably the most common basic means of animal communication with even the
most primitive animals reacting to odours given off by their own or other species.
• Animals may use scents to proclaim their readiness to mate, to mark out territorial boundaries,
to warn off intruders and predators or, in some cases, to attract prey. The most basic substance
for these purposes is strong smelling urine, but there are other and more refined methods of
producing lingering odours. These take the form of scent glands which may be situated in the
hooves, on the face or close to the anus or scrotum.
• Communication by scent can prove dangerous to certain animals such as the musk deer and
certain civets. Their scent is so strong and persistent that their glands are highly prized by the
makers of perfumes who use it as a stabiliser.
• Bees, wasps, ants, moths and other insects rely largely on upon pheromones as a means of
communication. Pheromones are chemical substances which may be secreted in urine, dung or
produced by special glands. They are usually given off by the female of the species to attract
males. For instance, the antennae of some male moths are so sensitive that they can detect the
presence of a female at a distance of more than two miles!
• Sharks have a particularly efficient sense of smell and some species can detect drops of blood
in the ocean at a range of about one mile.
• Snakes have the most unusual means of detecting scents. They use their tongue to pick up
scent particles in the air. Having flicked out its tongue the snake then returns it to a special
organ in the roof of the mouth which analyses the scents collected.

Dung is another readily available source of scent used by many animals to mark their
territory, with some species accumulating and regularly adding to large heaps of droppings
which act as territorial marker posts. The rabbit, vicuna, West African civet and hippopotamus
are just four examples of these.

2. Sound
• If the olfactory signs are the most common form of communication among animals the
acoustic signals must surely be the next in line, as animals of all kinds rely to a great extent on
their hearing ability in order to succeed and survive.
• Among mammals, generally speaking, small animals squeak and large ones rumble. The
reason for this is that the smaller the animal's head, the higher the frequency of sound it can
receive and transmit.
• In order to determine which direction a sound is coming from, the brain must distinguish
which ear it reaches first, and the time difference between them. The closer the animal's ears
are together, the shorter the wavelength, and therefore the higher the frequency, will be
necessary for good discrimination.

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• However, just to show that nothing in Nature can be taken for granted, some small-skulled
species of birds can communicate with ultra-low sounds. In the case of mammals it must also
be remembered that a large head and body allow space for bigger vocal equipment - so a
roaring gerbil would seem quite out of place.
• The acoustic communication signal most frequently heard by humans is, of course, bird song.
On a still summer evening the territorial song of the male blackbird will carry a considerable
distance.
• If we were to listen carefully we might also hear the chirping of crickets and grasshoppers
together with the rather similar sounds made by shrews, but many of the acoustic signals
transmitted by animals go unheard or unnoticed by humans.
• The largest of all animals, the whales, appear to be experts in the art of communication by
sound, with each whale of the same species having its own repertoire of 'songs' which it
repeats at intervals. Smaller whales such as the dolphins and porpoises also have a wide range
of sounds, although these tend to be the clicking kind rather than 'songs'. These sounds alter
quite noticeably when dolphins are hunting prey - being much more rapid and excited.
• Very small creatures must also communicate with others of their kind and a good example of
this is the acoustic signal of the death-watch beetle. Animals who live in a dense habitat such
as wood transmit information by a form of Morse code achieved by banging their heads
against the roof of their tunnel. This means of communication is clearly heard by humans.
• Frogs too are frequently heard by humans, and frogs and toads have developed their vocal
contact to a fine art by using bags of air as resonators. These may be situated in the mouth,
throat or on the side of the head and they are acoustically most effective. A species of
American tree frog provides an excellent example of the complex nature of communication
among animals.
• Although this frog gives a two-part call which humans hear in full the male frogs hear only
the first part of the call which warns them of the presence of a male intruder, while the
females hear only the second part which informs them of the presence of a potential mate!
• Grasshoppers and crickets create sound by 'fiddling' - a process which consists of rubbing the
hind legs over the ribs of the forewings. Male gorillas will beat their chest with cupped hands
to produce a sound which carries some distance. This action is thought to be a form of
communication although it could also be a means of releasing tension.
• Elephants communicate by a series of rumbling noises made in their throats and trunks,
although a cow elephant calling her calf will simply slap her ears against the side of her head.
Trumpeting is restricted to moments of extreme excitement or danger.

Kangaroos, hares and rabbits will thump their hind legs on the ground as a warning signal.
The rattlesnake gives a distinctly audible and sinister sounding warning by vibrating its bell-
shaped tail segments while other snakes, lizards and crocodilians will hiss loudly to warn off
intruders.
• The study of acoustic communication among animals is producing new discoveries each year
- leaving us to marvel at the complexity of Nature and the way in which life on Earth has
developed.
• 3. Sight
• Visual signals may take the form of gestures and display, facial grimaces, body posture or
mimicry. The extravagant display of the peacock or lyre bird can make the strutting of a wood
pigeon seem ineffective and yet each species has its own way of using visual communication
to the best advantage.
• The male rabbit will use the white underside of its tail to attract the attention of a female,
while the female may use the white of her tail as a visual signal for her young to follow when
she is leading them to the safety of the burrow.
• Dogs and wolves make use of body language, as do cats, monkeys and many other animals.
The attitude of the tail when two wolves meet will indicate which of the two is the superior.
The tail held between the legs is a submissive gesture while the tail raised confidently aloft
donates dominance. Horses will hold their ears or tail in certain positions to signal pleasure or
alarm. Dogs wag their tails as a sign of pleasure while the waving tail of a cat signals irritation
or anger.
• Apes and monkeys make considerable use of facial grimaces in order to express their feelings.
The frown of a rhesus monkey is a clear sign of unease, whereas the raised eyebrow and
fluttering eyelid denotes friendship or pleasure. Raised hackles in the Dog family help to
make the individual appear bigger when facing a possible adversary.
• At another level animals may communicate a simple message through the art of mimicry. The
roundels on the wings of the peacock butterfly look like large eyes to a potential predator. The

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harmless hoverfly resembles the wasp. Many caterpillars through their colouration take on the
appearance of poisonous or evil-tasting creatures in order to deter their enemies.
• With a little luck and a great deal of patience it is possible to observe and recognize visual
communication among animals such as birds or insects in the garden or dogs and cats in the
street.
• 4. Touch
• Although perhaps not as important or widespread as sound, scent or sight, a number of
animals make use of touch in order to communicate their feelings to other of their kind.
• Humans may shake hands, kiss or hug when meeting as a sign of respect or affection.
• Apes and monkeys will also hug and 'kiss' on meeting, although in certain species of monkey
the visiting or newly arrived monkey will place its hand in the mouth of the monkey it is
greeting. After a few moments the hand is withdrawn and the other monkey will place its hand
in the mouth of the visitor. This appears to be a sign of trust and goodwill between monkeys.
• Big cats tend to nuzzle each other, as do rhinos and many other animals. Elephants too use
touch as a means of close communication - interlinking trunks.
• It would take several lifetimes of study and a whole library of books to cover the fascinating
and often astounding study of animal communication and so we can only lightly scratch the
surface with a fact sheet of this kind. However, perhaps some of the topics covered by this
bulletin will fire the imagination of a number of young people who may become experts in
this field.
Humans can learn
Although humans do most of their communication through the use of words and obvious gestures,
much can be learned from observing the subtle communications of animals.
Body language, the tone of a person's voice and body odors can provide considerable amount of
information about the person's intentions, mood and even health. Often people react to these subtle
forms of communication without realizing that there has been some transfer of information.
Pheromones
Pheromones are chemicals released by living organisms that send information to other organisms of
the same species via scent. These pheromones are released in response to stress, alarm, danger, and
sexual fertility. They are released by both insects and mammals in many situations.
Insect Pheromones
• Alarm Pheromone
In response to danger or stress, ants release a chemical signal that is sensed by other ants nearby. In
response, these nearby ants travel towards the scent and work quickly in order to help their
endangered nest mate. Other insects, including honeybees, have these alarm signals that are released
via pheromones.
• Trail Pheromone
In addition to releasing scents in response to danger, ants also release a scent when they are returning
to their nest with food. This pheromone signals to other ants that there is food at the nest. As more
ants follow this scent, it is renewed in order to tell more ants. As food quantities decrease, ants release
a different scent that tells the others that there is no more food.
Sexual Attractants
Females most often release pheromones that attract males.
Mammal Pheromones
• Releaser Pheromone
Releaser pheromones bring forth specific behavior. Many mammals release pheromones that signal to
others that they are in their “territory.” This is seen in dogs urinating and also in many other
mammals. There are also other reasons for releaser pheromones. For example, female rabbits and
some other mammals release a scent that tells their young that they are ready to nurse. This scent then
causes their young to immediately begin nursing.
• Primer Pheromone
Primer pheromones cause a shift in the endocrine system of the animal receiving it. Many mammals,
including rats and mice, release pheromones that cause sexual behavior in the other sex. When at
peak fertility, females release a scent that attracts a male and causes him to become aroused and
exhibit mounting behavior on the female, who is also exhibiting lordosis. The pheromones are sensed
by the vomeronasal organ (VNO) in the brain. Another example of a primer pheromone is in rats.
Female rats mature faster when exposed to the scent of an adult male rat. Also, the scent of a male rat
of one strain can induce miscarriage in a female rat pregnant by a male of a different strain.
Information Pheromone
Information pheromones provide information about an animal’s identity. By smelling another animal
of the same species, they are able to tell what the animal ate last, if they are in heat, their level of
dominance, and their level of health.

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The Evolution of a Pheromone Signaling System:
From Molecules to Mating
Tens of millions of years ago, plethodontid salamanders evolved a bizarre system of chemical
communication in which courting males "vaccinate" females with a pheromone produced by a gland
on the male’s chin. In their subsequent history, plethodontids modified this pheromone delivery
system in a variety of ways, evolving other methods of pheromone delivery. This chemical
communication system – unique among vertebrates – has been retained by the diverse tribes of
plethodontids throughout their evolutionary radiation. We have established that the plethodontid
pheromone blend increases female sexual receptivity, indicated by faster time to insemination during
courtship. We have also determined both the amino acid sequences of individual pheromone
components and the nucleotide sequences of the genes that produce those components, establishing
that the pheromone is a blend of proteins from three unrelated families. The phylogenetic setting for
these discoveries offers an outstanding opportunity to investigate the evolution of the causal pathway
from gene to protein to behavior. This project seizes that opportunity and pursues it in a conceptual
framework that is generally applicable to the study of evolution at multiple levels of organization.
FROM MOLECULES TO MATING The opportunity presented by the plethodontid courtship
system is exceptional because of the direct path from gene to behavior. It is rare to connect a
particular gene to behavior, and especially to an important behavior. The important behavior in our
system is female sexual receptivity, a behavior at the crux of burgeoning research in the field of
evolutionary biology. Receptivity is a central issue – perhaps the central issue - in sexual selection,
sexual isolation, and speciation. By discovering the only vertebrate pheromone known to alter female
receptivity and establishing that it is a protein, we have opened an important window on evolutionary
pattern and process. The aim of this project is to look through that window at multiple levels: at the
level of the genome, at the level of expressed protein, at the level of neuroendocrine pathway and at
the level of female sexual receptivity.
EVOLUTION AT DIFFERENT LEVELS IN A FUNCTIONAL COMPLEX In this project we
tested for alternative evolutionary modes (stasis vs. rapid change) at multiple levels in a chemical
communication system. There is no one unifying construct that makes theory-based predictions at all
levels (e.g., from proteins to neural response). Sexual selection theory, for example, can address
expectations concerning a male signal and the female response, but offers little insight into structural
changes of a three-dimensional protein signal. We therefore used a variety of perspectives to make
predictions about evolutionary stasis or change.
These The six perspectives and their predictions are:
(1) A sexual selection perspective on communication systems predicts either stasis or rapid
evolution, depending on whether selection acts on the receiver.
(2) An empirical perspective based on observations of morphological and behavioral evolution
in Plethodon salamanders – predicts either stasis or rapid evolution, depending on which branch of
the phylogeny is under scrutiny.
(3) A neurophysiological perspective based on the pathways mediating female behavioral response
to the male signal – suggests that radical change in both the signal and receiver should co-occur in
one particular branch of the Plethodon phylogeny.
(4) A molecular perspective on the cytokine protein family to which one of the pheromone
signals belongs suggests that some parts of the pheromone signal should be static (e.g., binding
sites), but other parts of the molecule should be capable of rapid evolution.
(5) A molecular perspective based on recent case studies of proteins suggests that our pheromone
signals are candidates for rapid molecular coevolution.
(6) Preliminary analysis of pheromone gene sequences from four Plethodon species reveals rapid,
selection-driven evolution of some parts of one pheromone molecule. Taken together, these
perspectives motivate a research strategy in which we focus tests on particular branches of the
Plethodon phylogeny. We ask whether the same kinds of selection (stabilizing or diversifying) and
same evolutionary mode prevail at all levels of the signaling system.

EVOLUTION OF BEHAVIOR
One general view in the study of the evolution of behavior is that behaviors can have a genetic basis.
This is not to say that all behaviors are genetically based; indeed many behaviors are entirely
culturally transmitted or learned and may have little to do with genetics (why are you sitting in the
same seat?). For genetically influenced behaviors we can treat them as we would treat any other
genetically controlled trait of an organism: 1) if there are genetically based differences in a behavior,
and 2) these differences affect fitness then, 3) behaviors can evolve by natural selection.
Two examples of genetically based behaviors: cricket song. Different species of crickets have
different calling songs with different characteristics, e.g., inter chirp interval, pulse repetition rate, etc.
Hybrids between closely related species often exhibit songs with intermediate characteristics

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BIOTRANCE
ANIMAL BEHAVIOUR
(pulse repetition will be intermediate, inter pulse interval will be intermediate, etc.) a hypothetical
example with time on horizontal axis and each chirp = a group of vertical lines:

Another example (on a larger phylogenetic scale) is head scratching with the hind leg in amniotes
(reptiles, birds, mammals; those with an amniotic sac). Most reach the hind leg over the fore limb to
scratch the head; that birds and mammals do it suggests that this behavior has a genetically
programmed basis and has been inherited through much of higher vertebrate evolution.
Behavior is usually dissected into two components for analysis: Proximate causes/questions in
which one asks how the behavior is performed and ultimate causes/questions in which one asks why
the behavior is performed. Tinbergen has identified four questions to pose when analyzing a
behavior 1) what is the cause, 2) what is the development (ontogeny), 3) what is the current function
4) what is the phylogenetic history. A strict course on evolution focuses more on the latter two
questions (recall adaptation/preadaptation/exaptation discussion and the identification of current
utility vs. historical origin).
Herring gulls breed is large colonies on the ground and defend territories. Two separate calls used for
1) advertising nest site ("choking" call) and 2) as a territorial claim (the "oblique pose" and "long
call"). The Kittiwake also breeds in colonies but nests on vertical cliffs and its nest pad is its territory
and breeding site. In this species only one behavior serves both functions: "choking" behavior is both
defensive and part of mate recognition/pair formation. This is seen as an adaptive behavioral shift wit
respect to the nest location (steep cliff).
There are many behaviors that at first appearance do not seem "adaptive". Infanticide in lions was
first viewed as "aberrant" behavior by abnormal individuals because it was not "good for the species"
(male lions displace other males from groups of females and their offspring, and frequently kill the
cubs). It is true that killing infants is not, in the short term, an effective means of increasing
population numbers of a species. BUT, we now know (post W.D. Hamilton's 1963, 1964 papers on
inclusive fitness and kin selection and G. C. Williams book on Adaptation and Natural Selection) that
the more appropriate way to address such problems is to think about them in the context of whether
the behavior is good for the individual.
In analyzing infanticide from the perspective of gene thinking it is 1) not adaptive for a male lion to
invest reproductive effort in an individual with whom he shares no genes and 2) once the infant is
killed it is advantageous for the female to come into estrous and have more offspring with the new
male (this will increase her reproductive output over leaving with the displaced male, and not
benefiting from other advantages of group living: foraging, avoiding predation on young). Given the
situation for both male and female, the observed behaviors make sense in terms of propagating ones
genes.
The role of the gene (or genes!) as the unit that is relevant in the evolution play an important part in
two influential books in the mid 1970s Sociobiology by E. O. Wilson, and The Selfish Gene by
Richard Dawkins. To grossly oversimplify one of their main messages: "an organism is just DNAs
way of making more DNA"
If we take the case of bird migration we want to know how the bird navigates to the breeding location
(solar and magnetic cues during flight), how the bird knows when to begin migration (internal clocks
and changes in day length [physiological changes]). There is usually a high cost associated with
migration so we also want to know why birds do it since many die in the process (more time for
feeding, more available food). Individuals that do migrate must leave more offspring than those that
do not - again gene thinking helps account for why the behavior exists
Population genetic approaches to the evolution of traits rarely tell us why a phenotype affects fitness
in a particular way; the models usually look at whether fitness increases or not. The optimality
approach to the analysis of behavior attempts to builds models where different behaviors are treated
as the traits and asks which one of these behaviors might evolve. The approach generally ignores the
mechanics of underlying genetic basis of the behavior (i.e., its mendelian and transmission
genetics). Optimal models assume there is a genetic basis and treat each behavior as a haploid
(asexual) trait that is inherited intact.
While Gould and Lewontin (and many others) have criticized optimal models, the builders of optimal
model (e.g., John Maynard-Smith, Univ. Sussex) argue that the models do not assume that the
organisms are optimal (because there are constraints on evolution of traits), but by treating the
problem as an optimality issue, it can tell you what kinds of behaviors might evolve.
Two general type of optimal models: frequency independent models are designed independent of
what other strategies are doing, and seek to define the conditions which might influence behavior

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BIOTRANCE
ANIMAL BEHAVIOUR
(recall the "optimal foraging" model we described in the adaptation lecture where a bird assess,
quality, availability, distance to food items, etc.).
Frequency dependent models are ones where the strategy of one type depends on the strategies and
frequencies of other types in the population. The general approach is to look for Evolutionary Stable
Strategies (ESS) = a strategy that, if adopted by all, cannot be "invaded" by a mutant strategy. Here a
strategy = the behavior of an individual in a certain situation. These types of model apply nicely to
ritualized behaviors, distinct display behaviors which take the place of aggressive interactions.
Maynard-Smith's approach involves:
H = hawk who fights until the opponent retreats or will continue fighting injured with cost C
D = dove who displays but will retreat if the opponent escalates
V = payoff of winning an encounter
C = cost of losing an encounter
These values can be put into a payoff matrix:
In encounter with:
H D
Payoff to: H 1/2 (V-C) V
D 0 V/
2
H:H interaction = 1/2(V-C) because each individual hawk will win half of the time and lose half of
the time. In the D:D interaction each will win half of the time and retreat half of the time (retreat with
no cost). Which strategy is an ESS? Answer by asking if a strategy can invade. Can H invade a
population of D's?: Is payoff (D against D) > payoff (H against D)? i.e., is V/2 > V? Answer = NO, so
H can invade a population of D's.
Is H an ESS? Is payoff (H against H) > payoff (D against H)? i.e., is 1/2(V-C) > 0? Answer: it
depends on the values of V and C: if V > C then payoff to H will be positive and H is an ESS; if V <
C then payoff to H will be negative and neither D nor H will be favored (H will always invade a
population of D's until H's become so frequent that they encounter each other frequently. D can
invade a population of H's because H's tend to damage each other too much. In fact a population of all
H's with V<C would go extinct. Thus which behavior evolves depends on the nature of the
interactions.
One can imagine many other games and payoff matrices that could be built to model other
behaviors. The point of all this is to imagine the following: some species have ritualized displays that
appear "civil" in an anthropomorphic sense. Have these behaviors evolved through a stage where
hawks killed each other (C was high) to their current state where the cost C to engaging in a behavior
is considerably less? This question could be addressed by comparing the behaviors of related species
and applying the game theory approach.

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