Greg Detre
Monday, 20 May, 2002
Recent critical reviews of the
known cases of imitation in primates challenge all such claims and offer more
parsimonious explanations based on social facilitation and stimulus enhancement
(Whiten 1989).
Similarly, teaching has been restricted to humans, although in one case a
sign-language-trained chimpanzee, attempted to mould and influence the signing
performance of a younger companion (Fouts et al. 1982)
Chimpanzee mothers may
influence the development of nut cracking in three ways (excluding the very
wide-spread nut sharing) (Boesch & Boesch 1984):
1. stimulating
2. facilitating nut-cracking
3. active teaching
They
observed two cases in which the mother, noticing the infants� difficulties,
made a clear demonstration of how to solve them, e.g. Ricci�s 5-year old
daughter, Nina
All functions attributed to
human tutoring by Wood et al. (1976) seem to be performed by
chimpanzee mothers here. Recruitment, direction maintenance and marking
critical features are apparent in the stimulations, frustration control and
scaffolding in the facilitations as well as in teaching alongside with
demonstration.
Common ravens have a
reputation for being intelligent. They are ecologically highly adaptable, are
both neophobic and neophilic (Heinrich, 1988, 1995), long-term monogamous and
territorial for most of their life, but also flock.
pine-cone stripping of black
rats (Zohar
& Terkel 1996) or the opening of milk bottles by great tits (Sherry &
Galef 1990).
Stimulus enhancement
observation
of a model�s activity makes the location or object of the model�s behaviour
attractive for an observer (Thrope 1963, Galef 1988, Campell & Heyes in
press). Heyes (1994) defined stimulus enhancement as one-stimulus learning,
involving no association between the location or object and the reward.
Observational conditioning
Defined
as a kind of classical conditioning, where the observer associates the location
or object with the reward obtained by the model (Cook et al. 1985, Heyes 1994).
Both
stimulus enhancement + observational conditioning can increase the probability
of the observer learning an operant task.
Motor imitation
Motor
imitation is defined as learning the operant task directly through the
observation of the model�s behaviour (Heyes 1994, Zentall 1996). It is usually
considered the cognitively most demanding category of social learning (for the
observer), since it requires the translation of a visual input into a matching
motor output which may involve more complex central processing than other
mechanisms of social learning (Whiten & Byrne 1988, Heyes 1988).
Although a number of
studies� have focused on the imitative
ability of different primate species (e.g. Byrne 1995, Whiten et al 1996,
Bugnyar & Huber 1997), a few nonprimate mammals, such as rats (Heyes et al
1992, Heyes 1996), and several bird species, e.g. budgerigars (Galef et al
1986), grey parrots (Moore 1992), pigeons (Zentall et al 1996) and Japanese
quails (Akins & Zentall 1996), the conclusiveness of the evidence for
imitation learning is still debated (Visalberghi & Fragaszy 1990, Byrne
& Tomasello 1995, Heyes 1998, Gardner & Heyes in press). Studies on
�lower� mechanisms of social learning, e.g. enhancement or observational
condition, are rare.
Palameta & Lefebvre�s (1985) study of pigeons showed that observers were only
sufficiently motivated to peck through a paper lid on dish when they saw a
conspecific doing it, but not when they only saw the model either pecking or
eating. This suggests that the pairing of a demonstrator�s response with a
secondary reinforcer is very important.
Dawkins (1993), �Through our eyes only�:
Brunton and Macdonald used radiocollars to follow rats� movements on a farm
where a major extermination attempt had failed to kill more than a third of the
rat population
Galef paired rats in cages, showed the observer rat to
avoid/choose foods that harm/feed demonstrator rats
Caro & Hauser do not think that teaching depends on complex
intentionality or attribution of mental states, although intentionality plays
some role in some forms of teaching
Thorndike (1898) provided a clear definition of imitation �learning to
act from seeing it done�
but I
don't think that�s clear enough � it doesn't take into account behaviour that�s
somehow/implicitly influenced by watching someone else as opposed to an actual
attempt to reproduce it???
Tayler and Saayman (1973) report the amusing story of a captive dolphin that,
after repeatedly seeing a diver cleaning a viewing window, started cleaning the
window with a seagull feather while emitting sounds almost identical to those
of the diver�s air-demand valve and relaeasing a stream of bubbles from its
blowhole
Hayes and Hayes (1952) report training a chimpanzee to obey the command �Do
this� with an attempt to copy their own behaviour. They further report that
their chimpanzee appropriated a lipstick, stood on the washbasin and applied
the cosmetic to her mouth.
Byrne and Byrne (1991) report that mountain gorillas learn to process herbs
in specialised fashions that vary across local populations
Kawai (1965) reported that potato-washing of a juvenile female, Imo,
was soon adopted by most of the younger members of the group. There has been
much subsequent questioning of this finding. Green (1975) noticed that only the
members of the group that showed this behaviour were given potatoes by
provisioner so that the behaviour could easily have been caused by simple
reinforcement. Galef
(1990) looked at the time scale over which the novel behaviour
spread and found that the mean and median times for the acquisition of this
trait were two years, surely imitation would allow behaviour to spread almost
instantly through a population, as is suggested by the experimental studies of
imitation.
Fisher and Hinde (1949) proposed that the annoying habit that tits developed
in urban areas of opening milk bottles and drinking the first few centimetres
could have been developed independently in a few tits and then been acquired by
the population through observational learning.�
The theory was based on the observation that the behaviour seemed to
start in one area and then spread outwards. Sherry & Galef (1990) have
shown experimentally with black-capped chickadees that na� birds that had
observed a model opening cream tubs are more likely than non-observers to adopt
the behaviour themselves.
Spence (1937) was the first to propose stimulus enhancement
Dawson and Foss (1965) developed the �two-action test� that distinguishes
imitation from local enhancement.� The
paradigm is that a na� animal will watch a model obtaining food by one of two
action patterns
Fritz and Kotrschal (1999) used the two-action test to investigate social
learning in ravens
Gallese & Goldman (1998) report finding neurons in monkeys that respond to a
particular behaviour whether it is performed by the subject or by a monkey that
the subject is watching
song dialects passed down
generations � very reliably for the short song of the black-capped chickadees (Ficken and Popp,
1995)
opening
of milk bottle tops by blue tits (Hinde & Fisher 1952)
but Sherry &
Galef (1984) showed that such cultural transmission could occur
without imitation (i.e. without one bird actually seeing another open a bottle)
European blackbirds give alarm
calls to a stuffed owl or even a plastic bucket if they had heard their
parents� alarm calls while looking at them (Curio et al. 1978)
cultural
evolution can affect genetic evolution, e.g. the Japanese macaques who learnt
to wash their food ventured towards the sea, then started swimming and eating
sweaweed etc. (Bonner,
1980 or Wilson, 1985)
Menzel & Erber (1978) honey-bees learn rapidly and retain very well the
effects of even a single association between a colour and food reward
some
animals store food from a rich food source for later, either in a single
larder, or scattered in caches around their territory (Sherry 1985), e.g. the marsh tit
or Clark�s nutcracker
Tolman (1932) rats build up a mental picture/cognitive map of the
whole maze during exploration + learning
K�r (1927) on chimpanzees � showed that they can overcome
increasing hurdles to get their bananas
Pepperberg (1990, 1991) on the African grey parrot (Alex) � can count to six,
identify the quality (i.e. shape, colour) common to a group of diverse objects
Harlow (1949) � a monkey is faced with a pair of objects, one with a
reward, one without � it learns which is which � then a new pair of objects,
same rules � after 100 or so tests, the monkey simply lifts up one, and then
selects that one ever after if it had the reward
Warren (1965) all vertebrates except fish have have the ability to
form learning sets
Humphrey (1976) the complexities of social life, especially in the
primates, provided one of the most important selective forces behind the
emergence of thinking ability
Povinelli et al. (1992): experiment that suggested that chimpanzees have the
ability to understand the outcome of their own actions, the way that these
affect another individual, and also to put themselves effectively into that
individual�s place. two chimps on either side of the apparatus � two food trays
inside � the experimenter shows the informant chimp which handle to pull (I
think) to get the food, who has to then gesture the information to the operator
chimp. the operator chimp has to appreciate the significance of the informant�s
gestures and, after the swap, take on this gestural role � it does so
spontaneously, and using its own gestures which are not necessarily the same as
those it had seen used before by its partner
Premack & Woodruff (1978): chimp was shown a video of a human being using
objects well-known to the chimp, and in need of something (e.g. shivering with
cold with an unlit stove, or trying to open a locked door). then the chimp was
shown photographs of objects, one of which offered a solution (e.g. the burning
wick of the stove, a key)
Pfungst (1965) Clever Hans
elephants:
large brains, complex social life, longevity comparable with ours. perhaps significant
infrasonic (8-10Hz) communication between them (Poole et al. 1988), Payne (1999)
Gould (1981) described intelligence as the ability to deal flexibly
or creatively with problems
Gardner (1983) placed emphasis on symbolic processing as the hallmark
of intelligence
Byrne (1995) gives the following definition: �the term intelligence
should be restricted to that quality of flexibility that allows individuals to
find their own solutions to problems; genetical adaptations, by contrast, are
fixed and inflexible, however well tuned to their environments they are.�
Tool use has been discovered
in other animals, sea otters use stones to open shellfish, chimpanzees use
twigs to fish for insects, even adapting the shape of the twigs to make them
more suitable, and wooden hammers to open nuts. These behaviours are not innate
and, certainly in the case of chimpanzees are learnt by young from their
mothers (McGrew,
1998)
Heyes (1998) �theory of mind� = �an animal with a theory of mind
believes that mental states play a causal role in generating behavior and
infers the presence of mental states in others by observing their appearance
and behavior under various circumstances�
i.e.
�mental state attribution� (Cheney & Seyfarth 1990)
Dennett�s levels of �intentionality�
(see Byrne,
1991)
zero-order intentionality of
the eyed hawk-moth (recounted in Byrne 1991)
learnt, non-intentional but
intentional-seeming behaviour can be divided into two types: a) associative
learning, or b) as a product of inferences about observable features of the
situation rather than mental states (Heyes, 1993)
Gilgil the female baboon�s
grooming + meat-snatching (observation by Strum, cited as personal communication in Jolly 1985)
Hauser (2000) relates his observation of Tristan�s alarm call when
being chased by Borgia
Whiten and Byrne�s (1988) 253 accumulated reports along similar lines are
persuasive
Gallup (1969) mirror test: �knowledge of mental states in others
presupposes knowledge of mental states in oneself and, therefore, that
knowledge of self paves the way for an inferential knowledge of others�
apparently,
this ability to recognise their own images in mirrors has been replicated over
20 times, with a variety of species, but mainly primates
Reiss & Marino (2001) two bottle-nosed dolphins, Presley and Tab could
recognise themselves in the mirror
Povinelli (1998) they may simply have extended their motor
self-concept, rather than have developed a psychological one � the evidence
that chimpanzees are unable to imagine someone else�s perspective is quite
strong
Cheney & Seyfarth (1980, 1990) found that vervet monkeys give alarm calls on seeing a
predator even if other monkeys have already seen it
de Waal chimpanzees do not appear to trust the reassurance gestures of their
former opponents unless such gestures are accompanied by a mutual gaze
Premack & Woodruff (1978) �Does the chimpanzee have a theory of mind?�
Wimmer & Perner (1983) devised the Sally-Anne false belief test for young
children
Baron-Cohen et. al. (1985) autistic children fail to make an important cognitive
leap that usually takes place around age four when children learn that others
may have different beliefs about the world from their own
Bloom and German (2000) false belief test places high cognitive and linguistic
demands on infants and animals, far beyond the possession of a theory of mind
Donald Davidson (1975) from Dennett (1995) �claimed that only creatures with the concepts of
truth and falsehood can properly be said to have beliefs, and since these are
meta-linguistic concepts � only language-using animals such as human beings can
have beliefs�
Dennett (1995) employs a �maximally permissive understanding of the
term� �belief� � �even amoebas - like thermostats - have beliefs�� cf also McCarthy (1979)
Halperin (1995) Siamese fighting fish as candidates for having beliefs
(intentional interpretation), but neural-net-like model of their control
systems that seems to lack any components with the features beliefs are often
supposed to have
Byrne (1995) great apes definitely do have a theory of mind, but
that the case for monkeys seems especially weak, and little better for dolphins
or other non-primates
Bright (1984) �Animal language�
Payne (1995) recordings of humpbacks in Bermuda
Tyack (1986) used playbacks of the social calls, which brought the
singing whales to home straight in on the loudspeaker at 12km/hour � considered
that the song may be used as a spacing mechanism for courting males
Norris (1991) studied the social structure and behaviour of groups
of Hawaiian spinner dolphins
Bastian (1964) conducted a famous experiment with two dolphins, able
to hear but not see each other
John Lilly probed dolphins� propensity to mimic by getting them
to follow a count up to ten and recognisably say simple English words
Fisher & Ford (1983) sounds of killer whale pods are very stable, comprised
of about 12 distinct, stereotyped calls, that they use to communicate when
spread over a couple of miles
Hockett (1960, 1966) devised a set of 13 linguistic universals:
semanticity
and arbitrariness, productivity, interchangeability, specialization,
displacement, cultural transmission, duality
of patterning; auditory-vocal
channel; broadcast transmission
and directional reception; discreteness;
rapid fading):
Gardner & Gardner (1989) Washoe
by the
age of five, she had mastered 133 signs. She had been spontaneously combining
signs after learning only 8 or 10
Terrace (1979) Nim Chimpsky, less successful, rewarded with approval
rather than food, used symbols in the absence of their referents, but not novel
generation
Fouts et al. (1989) no sign language was used by humans in Loulis�
presence, yet he was still able to learn 51 signs just from the other
chimpanzees
Kanzi (Savage-Rumbaugh 1991)
Pepperberg (1991) Alex the grey parrot
our ethical anthropocentricism
may be regarded in the same light that slavery is now (Cavalieri & Singer 1993)
Wilson (1975) �Communication occurs when one individual�s actions
provide a signal that changes the behaviour of another individual�
Shannon (1948)
Wiley (1983) �noise� in terms of attenuation, extraneous
environmental distractions and any deficiencies in the signaller�s encoding or
receiver�s decoding equipment
rufous-sided towhees (Richards 1981)
�alerting signal� an initial burst of loud sound of unvarying pitch preceding
the more complicated warbling of the song which follows
Dawkins & Krebs (1978) case for �manipulation�, where every communication
balances the selfish attempt of the signaller to increase its fitness, against
the selfish attempt of the receiver to increase its fitness
Hamilton (1964) presented this as a 2x2 matrix, ranging from mutuality
(where both signaller and receiver benefit), deceit, eavesdropping and �spite�
(where both suffer from the communication).
Rohwer�s (1977) experiment with black patches on Harris� sparrows
exemplifies how the costs to the signaller of being probed and exposed may be
high
young male elephant seals (Le Boeuf 1974)
who have evolved to look like females in order to sneak past the bull elephant
into his harem suffer costs from probing by the female receivers, who emit loud
screams during copulation
red deer (Clutton-Brock & Albon 1979)
emphasis placed by ethologists
on handicapping signals (Zahavi 1975) has since been undermined (Maynard Smith
1976)
Blumberg & Alberts (1992) caution distinguishing functions and effects (e.g.
humans babies with respiratory distress syndrome that paediatricians listen
for)
vs similar laryngeal braking
in the Norway rats (Blumberg & Alberts 1990), and the mother�s
acute auditory sensitivity in this range, they assume that the ultrasonic
squeaks produced are some sort of �distress call�
different modalities:
birds
may have very conspicuous ultra-violet markers (Burkhardt 1989), which they use as
a private channel of communication
or
snakes see infrared radiation from warmblooded prey (Newman & Hartline 1982)
Von Frisch (1967) showed that bees can use the pattern of polarisation
of light to locate the sun�s position even when it�s obscured (with a small
dorsal region of the retina with a row of analysers each maximally sensitive to
a different e-vector direction)
smell,
e.g. mice � a pregnant female mouse will abort her entire litter if she smells
a strange male mouse near (because they eat newly born mice that aren't theirs)
(the �Bruce effect�)
odour
cues associated with the MHC affect behavioural (i.e. who a mouse nests with,
chooses as a mate (Lenington, 1994)) as well as cellular
recognition
may be
the case with human females rating the smell of t-shirts � reported that
pleasanter� from men most different from
them in the MHC (Wedekind et al. 1995), and that the disimilar ones reminded
them most of their former mates
deaf turkey hens kill most of
their chicks because they never receive the auditory sign-stimulus for parental
behaviour (Schledit et al. 1960)
Tinbergen (1951) pointed to false positives as one of the most
conspicuous characteristics of innate behaviour
e.g.
male sticklebacks (that normally respond aggressively to the red colouration of
rival male sticklebacks) also display to a red mail van visible through the
windows of the aquarium
Lettvin (1959) six different sorts of frog ganglion cells
Arak and Enquist (1993) supernormal stimuli emerge spontaneously in
computational models that rely on a few key sign stimuli
males sporting super-normal
plumage may be favoured, leading to an evolutionary exaggeration of both male
plumage and female preference� (Fisher, 1958)
Davies and Brooke (1989) showed that spotted flycatchers and reed buntings
(which currently don't get parasitised) were very good discriminators of model
cuckoo eggs, whereas meadow pipits were much less discriminating but don't get
parasitised because their diet/nest site aren't appropriate for cuckoos
sensory
systems are adapted to pick out �information-rich� parts of the environment (Barlow 1972)
� postulated grandmother cells at the top of the hierarchy
separate pathways in mammals
for movement + form (Zeki 1993)
e.g. neurons in the temporal
lobe of rhesus monkeys that fire up to 20 times more rapidly when looking at a
face (monkey or human) than for lines, gratings or non-face complex objects (Baylis et al.
1985)
we would say that the scent
trail is a signal used for communication by the ants, but although the snake
undoubtedly obtains information from it, we wouldn't want to say that the ants
communicated with it (Burkhardt, 1970)
Altmann (1962): social communication = a �process by which the
behaviour of an individual affects the behaviour of others�
Hinde + Rowell (1962): classified as visual signals only those that were
likely to have evolved for this purpose
van Rhijn (1980) saw that younger jays tend to give way to more
powerful older members, even with only a minimal �signal� (e.g. displacing
younger ones from food just by looking at them) once the rank order had been
established (although they would escalate into a recognisable threat display if
the minimal signals were ineffective)
same with dogs + coyotes,
bowing + wagging their tails (Darwin 1872, �The expression of the emotions in
man and animals�)
chimpanzees touch and kiss
each other�s hands in gestures of reconciliation (de Waal 1996)
Payne & McVay (1971) humpback whales may communicate over distances of
hundreds+ miles
hyenas smear grass stems with
past from their sub-caudal scent glands and deposit faeces at latrines, either
all round, or just in strategic places if the borders are too big (Mills &
Gorman 1987)
the
extent to which signaller and receiver�s interests coincide is an important
factor in how conspicuous the signals are (Dawkins & Krebs 1978), e.g.:
red
deer (see below)
gazelles
stotting (see below)
on
the other hand, cooperating animals communicate with �conspiratorial whispers�
(Krebs &
Dawkins 1984)
conflict
between signaller + receiver leads to selection pressure for signals to be
�honest�, i.e. proof against cheating/bluffing (Zahavi 1987)
red
deer (Clutton-Brock
& Albon 1979)
gazelles
(Fitzgibbon
& Fanshawe 1988)
skylarks
sing when being chased by merlins (Cresswell 1994)
young
canaries� begging signal (wide open mouth) � their mouths� red lining is due to
the suffusion of blood � parents feed the reddest mouths, as their means of
deciding whose need is greatest (Kilner & Johnstone 1997) � after being fed,
the blood is diverted to the gut, and the mouth gets paler
Ristau (1991) argues that plovers giving a broken wing display
monitor the behaviour of the predator (e.g. fox) and adjust their behaviour
according to whether the fox is following them or still heading for the nest
Munn (1986) describes two species of insect-eating birds that
appeared to deceive their flockmates by falsely giving alarm calls and so
gaining unrestricted access to food
Byrne & Whiten (1988) consider possibly deliberate baboon behaviour � although
anecdotal, they argue there are enough examples to support the idea that some
animals can be deliberately dishonest
Gould (1975) summarised + proved von Frisch�s (1967) conclusions
Cheyney & Seyfarth (1980, 1990) vervet monkeys (Africa),� elegant + group-living
evolutionarily stable strategy
(Maynard Smith
1982) � an ESS is a strategy that, if adopted by most members of the
population, cannot be bettered by any other strategy (one new mutant strain)
domestic fowl (Schjederup-Ebbe
1935) linear hierarchy � lots of fighting to begin, but once
established, subordinates almost always defer to a more dominant bird
the black bibs of male house
sparrows � if a weak animal has a dominant badge, it gets constantly challenged
by other dominant individuals (Moller 1987), and so pays a cost for its
deception (which may be enough to keep the number of cheats pretty small)
R. Dawkins (1976) �The Selfish Gene�, green beard, memes
Dennett (1987) �The intentional stance�
Hamilton, W. D. (1964) �The genetic evolution of social behavior�
Axelrod & Hamilton�s (1981, 1984) theoretical work on the Prisoners� Dilemma
Wilkinson�s (1984) observations of regurgitation of blood by vampire bats
lionesses are far more
effective hunting zebras together than alone (Caraco & Wolf 1975)
Jarvis (1994) considers two species of mole-rat
Holmes and Sherman (1982) showed that sibling recognition in Belding�s ground
squirrels is heavily skewed by those they have been reared with
Yamazaki et al. (1976) mice preferred to mate with mice that differed at the
MHC even when genetic background was controlled
Brown & Eklund (1994) MHC similarity outweighs the effect of the entire
remaining genetic background
Belding�s ground squirrels (Holmes &
Sherman 1982)
in the sweat bee, the
willingness of the guard bee to let in other bees was directly correlated with
their relatedness (Greenberg 1979)
Emlen (1990) reports that 45% of helping in the white-fronted
bee-eater is directed to rearing full siblings, and that on average r=0.33 between
helpers and nestlings
Grafen (1990) argues convincingly that in the majority of cases
where kin recognition has been reported, the ability to do so has arisen as a
result of some other function and so should not be termed kin recognition as
such; indeed, in many of these cases he argues that the demonstrated ability to
recognise kin is not selectively advantageous or even of any functional value
at all.
Manning et al. (1992) - house mice - communal nests, nurse other�s pups
indiscriminately; kinship theory - should form nests with relatives to minimise
exploitation and increase inclusive fitness? Females prefer communal nesting
partners that share allelic forms of mhc genes. Grafen�s (1992) criticisms
Blaustein suggests four possible
mechanisms for recognition:
based on spatial distance
based on familiarity, prior association
based on phenotype matching
based on action of recognition alleles
Allee et al. (1938) showed that water fleas cannot survive in alkaline
water, but the respired CO2 of a large group could be sufficient to
bring down the alkalinity
adult meerkats (socially
living mongooses) take turns �baby-sitting� the nest-hold (watchign for
predators from a high look-out point) while the others are away foraging (McDonald 1986)
group living mongoose,
satiated adult adopts raised sentry position while the other group members
forage for food, watchman�s song (Benekoff, 1997)
naked mole rats (Jarvis 1981)
average relatedness 0.81 (monogamy plus in-breeding), wouldn't survive on their
own or in pairs, tips the cost-benefit-relatedness analysis in favour of
self-sacrificing, life-long worker sterility
Damaraland mole rat (Jarvis et al.
1994)
Florida scrub jays (Woolfendon &
Fitzpatrick 1984): helpers gain by having more siblings, they tend
to take over their parents� territory in the future (at the cost of deferring
reproduction for a year or so, but nest sites are highly in demand)
snapping shrimp (Duffy 1996)
lives in sponges on tropical reefs, in colonies of <300, but only one
reproductive female (the �queen�). like termites (Wilson, 1971) and mole rats, the
shrimps are diploid and the �workers� are full sibs. food seems to be abundant,
so the workers are mainly for defence, because sponge �homes� are sought by
various (larger) species on the crowded reef
multiple mating is not in the
interests of males, e.g. male dragonflies� penises have elaborate hooks to try
and scoop out the sperm from other mates stored by the female when mating (Waage 1979)
huge male elephant seals
dominate stretches of beach and fight over the females � in some seasons, 4% of
the males are responsible for 85% of the matings (Le Boeuf & Peterson 1969)
no complete agreement on why
primate organisation is so diverse (Ridley 1986)
Rowell (1974) critised the way the notion of dominance hierarchies
has been used: dominance may be more important in captivity than in the wild
but pretty linear hierachy in
Barbary macaques (Deag 1977)
Kummer (1969) transplanted a few yellow baboon females into a
hamadryas troop, which were quickly herded into harems � they didn't respond to
the male�s stare, and fled, but were chased back into the harem, and learnt
their lesson � thus, at least some of the variation in social organisation
between primate species is cultural rather than genetic
Zahavi (1974, 1995) showed that Arabian babblers would pretend to feed the
young in the nest and if there were no other adults near-bye would then eat the
food themselves, suggesting that there must be benefits in being thought of as
a good helper or the animals
Bateson
(1982, 1988), himself, found that quail prefer the company of first cousins to
brothers or third cousins and that birds mated to first cousins breed rather
earlier.
In mice the MHC genes have
been shown to alter behaviour in three contexts. It has been shown that mice
prefer to mate with individuals differing from themselves at this group of loci
(Yamazaki et
al., 1976). Females are more likely to nest communally with females
that are the same at the MHC (kin selection), a pregnant female will often
abort a litter or engage a �pregnancy lock� if she smells a strange male (Lenington, 1994).
This is clear evidence of individual recognition and a corresponding active
change in behaviour. More information will be discovered about the genetic
influences of kin recognition as genetic understanding and technology improves.
Porter et al. (1978) showed that spiny mice prefer to associate with
siblings rather than non-siblings but that the discrimination was based on the
mice an individual had been housed with and was therefore altered by
familiarity
Grafen (1990) defines recognition as �recognition by genetic
similarity detection.�
Wilkinson (1984) investigated food sharing amongst vampire bats
Grinnel, Packer and Pusey
(1995) suggest that there are three possible routes through which cooperation
may develop:
1. kin selection (Hamilton 1964) is the theory that the
inclusive fitness effects of cooperation can outweigh the temptation to defect
because the payoff for cooperation exceeds that of defection.
2. reciprocity (Trivers, 1971; Axelrod and Hamilton, 1981)
a theory that involves looking at repeated interactions with reference to
previous behaviour, reciprocate cooperation and exclude defection
3. mutualism (Maynard Smith, 1983; Lima, 1989) when it is
considered that there is no temptation to defect because the payoff from
cooperating is always higher no matter what the behaviour of the opponent. The
first two theories involve conditional cooperation, mutualism is unconditional
because the individual will always cooperate.
Nowak & Sigmund (1993) win-stay, lose-shift (Pavlov)
One of the most commonly cited
examples of cooperation amongst animals is the lion. They rear cubs, defend
territories and hunt together. Legge (1996) describes them as �an emblem of
cooperation.�
Levy (1992)
R. Dawkins (1986) �The Blind Watchmaker�
Paley (1802), �Natural theology�, Paley�s watch metaphor
Reynolds (1986) boids (separation, alignment, cohesion)
Mitchell (1998), common GA methods: �populations of
chromosomes; selection according to fitness; crossover to produce new
offspring; and random mutation of new offspring�, GA = biologically-inspired
�parallel population-based search [for solutions] with stochastic selection of
many invidividuals, stochastic crossover and mutation�
Holland (1975) early GAs
Hillis (1990) coevolution in sorting GAs
Yaeger (1992) even recounts how he hadn�t even realised
that he�d forgotten to switch mutation on for the first weeks of his PolyWorld
investigation
Anastasoff (1999) evolving, dynamic level of mutation
Rasmussen (1991), 'Aspects of Information, Life Reality, and Physics'.
1.)
A universal computer is indeed universal and can emulate any process (Turing)
2.)��� The essence of life is a process (von
Neumann)
3.)��� There exist criteria by which we are able to
distinguish living from non-living things
Accepting
(1), (2) and (3) implies the possibility of life in a computer
Conway�s (1970) game of life
Pinker (1997)
Griffin (1992) �Animal minds�
Nagel:
(1974) �What is it like to be a bat?�
(1979)
�Panpsychism�
Dennett (1991) �Consciousness explained�
M. S. Dawkins (1993), �Through our eyes only�
Jaynes (1976) �Origin of
consciousness in the breakdown of the bicameral mind�
�auto-pilot� (Baars 1988, Weiskrantz 1995
(the �British Weather Conversation Syndrome�))
Greenfield (2001)
Wittgenstein (1953), �Philosophical investigations�
Tinbergen (1963) �On aims and methods of Ethology�) � causation,
survival value, ontogeny and evolution
based
on Huxley�s
(1942) tri-partite distinction
�Behavior is all
observable or otherwise measurable muscular and secretory responses (or lack
thereof) and related phenomena in response to changes in an animal's internal
or external environment.� (Grier and
Burk, 1992)
M. S. Dawkins (1989) �The future of ethology: How many legs are we standing
on?�
Hailman�s (1977) partitioning individual/population and cause/origin
Mayr (1963) for example stresses the dichotomy between functional
and evolutionary biology, using the terms �proximate cause� and �how� questions
in relation to functional biology and �ultimate cause� and �why� questions in
relation to evolutionary biology
Dewsbury (1992) attacks the proximate-ultimate distinction
Gould & Lewontin (1979), �The Spandrels of San Marco and the Panglossian
Paradigm: A Critique of the Adaptationist Programme�
�The
design is so elaborate, harmonious, and purposeful that we are tempted to view
it as the starting point of any analysis, as the cause in some sense of the
surrounding architecture. But this would invert the proper path of analysis.
The system begins with an architectural constraint: the necessary four
spandrels and their tapering triangular form�
Dr.
Pangloss: "Things cannot be other than they are... Everything is made for
the best purpose. Our noses were made to carry spectacles, so we have
spectacles. Legs were clearly intended for breeches, and we wear them."
Harner (1977) proposed that Aztec human sacrifice arose as a
solution to chronic shortage of meat (limbs of victims were often consumed, but
only by people of high status)
the eskimo face, once depicted
as "cold engineered" (Coon, et al., 1950), becomes an adaptation to
generate and withstand large masticatory forces (Shea, 1977).�
this nonadaptive hypothesis
(about T-rex�s short forelimbs) can be tested by conventional allometric
methods (Gould,
1974, in general; Lande, 1978, on limb reduction)