Right to truth is the right, in the case of grave violations of human rights, for the victims and their families or societies to have access to the truth of what happened. The right to truth is closely related to, but distinct from, the state obligation to investigate and prosecute serious state violations of human rights. Right to truth is a form of victims' rights; it is especially relevant to transitional justice in dealing with past abuses of human rights. In 2006, Yasmin Naqvi concluded that the right to truth "stands
somewhere on the threshold of a legal norm and a narrative device ...
somewhere above a good argument and somewhere below a clear legal rule".
According to Patricia Naftali, the right to truth remains elusive
because it is a concept with different definitions (sometimes
contradictory), which is deployed in support of a variety of human
rights claims.
Take steps to avert re-occurrence of the violation
Change national laws
Institute measures to improve compliance with international human rights instruments
Construct memorials to commemorate the human rights violation
United Nations Human Rights Committee
The first case that articulated a right to truth in international human rights jurisprudence was a forced disappearance case, Quinteros v. Uruguay (1983); the UN Human Rights Committee determined that, according to the International Covenant on Civil and Political Rights,
the mother of the victim had "the right to know what has happened to
her daughter. In these respects, she too is a victim of the violations
of the Covenant suffered by her daughter in particular, of article 7
[ICCPR]". In Saadoun v. Algeria (2003), regarding a man who was forcibly disappeared during the Algerian Civil War,
the Committee determined that failure to investigate gave rise to a new
violation of the ICCPR. In this case, Algeria had proclaimed an amnesty
for crimes committed during the "national tragedy".
the right to the truth is subsumed
in the right of the victim or his next of kin to obtain clarification of
the facts relating to the violations and the corresponding
responsibilities from the competent State organs, through the
investigation and prosecution established in Articles 8 and 25 of the
Convention.
European Court of Human Rights
There is also case law of the European Court of Human Rights relevant to right to truth. In Cyprus v. Turkey (2001), the ECtHR ruled against Turkey in the case of Greek Cypriots
who had been last seen in the custody of Turkish troops. The anguish of
surviving relatives constituted a "continuing violation of Article 3 of the European Convention on Human Rights (ECHR) with respect to the relatives of the Greek-Cypriot missing persons." In El-Masri v. Macedonia
(2012), the ECtHR established that North Macedonia had violated the
Convention in allowing El-Masri to be taken into US custody during extraordinary rendition.
The court noted that Macedonian authorities had "deprived the applicant
of being informed of what had happened, including of getting an
accurate account of the suffering he had allegedly endured and the role
of those responsible for his alleged ordeal" as well as hidden this
information from the public at large. According to law professor Arianna Vedaschi,
"the decision given in El-Masri showed innovative legal reasoning and a
wholly innovative attitude of the judges towards the far-reaching
enforcement of the right to the truth". In Janowiec and Others v. Russia (2013), the court found no violation of the convention regarding Russian investigations into the 1940 Katyn massacre, but this ruling was on the principle of non-retroactivity because the massacre happened before the ECHR was drafted.
Legal scholar James A. Sweeney criticized the ECtHR's approach to right-to-truth cases:
the ECtHR’s 'underlying values'
test could have led the way in promoting internationally the notion that
present-day denial or obstruction of the quest for truth about the
gravest pre-ratification human rights abuses may amount, in itself, to a
contemporary human rights violation. Such an approach does not apply
each human rights treaty retroactively, nor does it convert every
historical human rights abuse into a 'continuing violation', but it
establishes exceptional circumstances in which denying the right to
truth about historical human rights abuses is constitutive of a fresh
violation within the temporal jurisdiction of the relevant enforcement
body.
According to legal scholar Agostina Latino, the right to truth related to the Armenian genocide extends beyond Armenian genocide survivors to their descendants as well as Armenians at large. Latino states that, as the successor to the Ottoman government that committed the genocide, the Turkish government's ongoing Armenian genocide denial violates their right to truth. For example, there are monuments and streets named after the perpetrators, but not the victims.
The Inter-American Court and some theorists have suggested that truth-telling may be a form of partial reparations to victims of human rights abuses. Right to truth is related to the fight against impunity as establishing the truth about a past event is the first step in holding perpetrators accountable.
Right to Truth Day
Since 2010, the UN has commemorated International Day for the Right
to the Truth Concerning Gross Human Rights Violations and for the
Dignity of Victims, or Right to Truth Day, on 24 March, the anniversary
of the murder of El Salvador archbishop Óscar Arnulfo Romero.
The co-operative behaviour of social insects like the honey bee can be explained by kin selection.
Kin selection is a process whereby natural selection favours a trait due to its positive effects on the reproductive success of an organism's relatives, even when at a cost to the organism's own survival and reproduction. Kin selection can lead to the evolution of altruistic behaviour. It is related to inclusive fitness,
which combines the number of offspring produced with the number an
individual can ensure the production of by supporting others (weighted
by the relatedness between individuals). A broader definition of kin
selection includes selection acting on interactions between individuals
who share a gene of interest even if the gene is not shared due to
common ancestry.
Charles Darwin discussed the concept of kin selection in his 1859 book, On the Origin of Species, where he reflected on the puzzle of sterile social insects, such as honey bees,
which leave reproduction to their mothers, arguing that a selection
benefit to related organisms (the same "stock") would allow the
evolution of a trait that confers the benefit but destroys an individual
at the same time. J.B.S. Haldane
in 1955 briefly alluded to the principle in limited circumstances
(Haldane famously joked that he would willingly die for two brothers or
eight cousins), and R.A. Fisher mentioned a similar principle even more briefly in 1930. However, it was not until 1964 that W.D. Hamilton generalised the concept and developed it mathematically (resulting in Hamilton's rule) that it began to be widely accepted. The mathematical treatment was made more elegant in 1970 due to advances made by George R. Price. The term "kin selection" was first used by John Maynard Smith in 1964.
According to Hamilton's rule, kin selection causes genes to
increase in frequency when the genetic relatedness of a recipient to an
actor multiplied by the benefit to the recipient is greater than the
reproductive cost to the actor. Hamilton proposed two mechanisms for kin selection. First, kin recognition
allows individuals to be able to identify their relatives. Second, in
viscous populations, populations in which the movement of organisms from
their place of birth is relatively slow, local interactions tend to be
among relatives by default. The viscous population mechanism makes kin
selection and social cooperation possible in the absence of kin
recognition. In this case, nurture kinship,
the interaction between related individuals, simply as a result of
living in each other's proximity, is sufficient for kin selection, given
reasonable assumptions about population dispersal rates. Kin selection
is not the same thing as group selection, where natural selection is believed to act on the group as a whole.
In humans, altruism is both more likely and on a larger scale
with kin than with unrelated individuals; for example, humans give
presents according to how closely related they are to the recipient. In
other species, vervet monkeys use allomothering,
where related females such as older sisters or grandmothers often care
for young, according to their relatedness. The social shrimp Synalpheus regalis protects juveniles within highly related colonies.
Historical overview
Charles Darwin wrote that selection could be applied to the family as well as to the individual.
Charles Darwin was the first to discuss the concept of kin selection (without using that term). In On the Origin of Species, he wrote about the conundrum represented by altruistic sterile social insects 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, and may thus gain the desired end. Breeders of cattle
wish the flesh and fat to be well marbled together. An animal thus
characterised has been slaughtered, but the breeder has gone with
confidence to the same stock and has succeeded.
— Darwin
In this passage "the family" and "stock" stand for a kin group. These
passages and others by Darwin about kin selection are highlighted in D.J. Futuyma's textbook of reference Evolutionary Biology and in E. O. Wilson's Sociobiology.
Kin selection was briefly referred to by R.A. Fisher in 1930 and J.B.S. Haldane in 1932 and 1955. J.B.S. Haldane grasped the basic quantities in kin selection, famously
writing "I would lay down my life for two brothers or eight cousins". Haldane's remark alluded to the fact that if an individual loses its
life to save two siblings, four nephews, or eight cousins, it is a "fair
deal" in evolutionary terms, as siblings are on average 50% identical
by descent, nephews 25%, and cousins 12.5% (in a diploid population that is randomly mating and previously outbred).
But Haldane also joked that he would truly die only to save more than a
single identical twin of his or more than two full siblings. In 1955 he clarified:
Let us suppose that you carry a
rare gene that affects your behaviour so that you jump into a flooded
river and save a child, but you have one chance in ten of being drowned,
while I do not possess the gene, and stand on the bank and watch the
child drown. If the child's your own child or your brother or sister,
there is an even chance that this child will also have this gene, so
five genes will be saved in children for one lost in an adult. If you
save a grandchild or a nephew, the advantage is only two and a half to
one. If you only save a first cousin, the effect is very slight. If you
try to save your first cousin once removed the population is more likely
to lose this valuable gene than to gain it. … It is clear that genes
making for conduct of this kind would only have a chance of spreading in
rather small populations when most of the children were fairly near
relatives of the man who risked his life.
W. D. Hamilton, in 1963 and especially in 1964 generalised the concept and developed it mathematically, showing that it holds for genes even when they are not rare, deriving Hamilton's rule
and defining a new quantity known as an individual's inclusive
fitness. He is widely credited as the founder of the field of social
evolution. A more elegant mathematical treatment was made possible by George Price in 1970.
The evolutionary biologist John Maynard Smith used the term "kin selection" in 1964.
John Maynard Smith may have coined the actual term "kin selection" in 1964:
These processes I will call kin
selection and group selection respectively. Kin selection has been
discussed by Haldane and by Hamilton. … By kin selection I mean the
evolution of characteristics which favour the survival of close
relatives of the affected individual, by processes which do not require
any discontinuities in the population breeding structure.
Kin selection causes changes in gene
frequency across generations, driven by interactions between related
individuals. This dynamic forms the conceptual basis of the theory of sociobiology. Some cases of evolution by natural selection can only be understood by considering how biological relatives influence each other's fitness. Under natural selection, a gene encoding a trait that enhances the fitness of each individual carrying it should increase in frequency within the population;
and conversely, a gene that lowers the individual fitness of its
carriers should be eliminated. However, a hypothetical gene that prompts
behaviour which enhances the fitness of relatives but lowers that of
the individual displaying the behaviour, may nonetheless increase in
frequency, because relatives often carry the same gene. According to
this principle, the enhanced fitness of relatives can at times more than
compensate for the fitness loss incurred by the individuals displaying
the behaviour, making kin selection possible. This is a special case of a
more general model, "inclusive fitness". This analysis has been challenged, Wilson writing that "the foundations of the general theory of inclusive
fitness based on the theory of kin selection have crumbled" and that he now relies instead on the theory of eusociality and "gene-culture co-evolution" for the underlying mechanics of sociobiology.
Inclusive fitness theory is still generally accepted however, as
demonstrated by the publication of a rebuttal to Wilson's claims in Nature from over a hundred researchers.
Kin selection is contrasted with group selection,
according to which a genetic trait can become prevalent within a group
because it benefits the group as a whole, regardless of any benefit to
individual organisms. All known forms of group selection conform to the
principle that an individual behaviour can be evolutionarily successful
only if the genes responsible for this behaviour conform to Hamilton's
Rule, and hence, on balance and in the aggregate, benefit from the
behaviour.
Hamilton's rule
"Hamilton's rule" redirects here. For the physical principle, see Hamilton's principle.
Formally, genes for a particular behavior should increase in frequency when
where
r = the genetic relatedness of the recipient to the
actor, often defined as the probability that a gene picked randomly from
each at the same locus is identical by descent.
B = the additional reproductive benefit gained by the recipient of the altruistic act,
C = the reproductive cost to the individual performing the act.
This inequality is known as Hamilton's rule after W. D. Hamilton who in 1964 published the first formal quantitative treatment of kin selection.
A 2014 review of many lines of evidence for Hamilton's rule found
that its predictions were confirmed in a wide variety of social
behaviours across a broad phylogenetic range of birds, mammals and
insects, in each case comparing social and non-social taxa. Among the experimental findings, a 2010 study used a wild population of red squirrels
in Yukon, Canada. Surrogate mothers adopted related orphaned squirrel
pups but not unrelated orphans. The cost of adoption was calculated by
measuring a decrease in the survival probability of the entire litter
after increasing the litter by one pup, while benefit was measured as
the increased chance of survival of the orphan. The degree of
relatedness of the orphan and surrogate mother for adoption to occur
depended on the number of pups the surrogate mother already had in her
nest, as this affected the cost of adoption. Females always adopted
orphans when rB was greater than C, but never adopted when rB was less than C, supporting Hamilton's rule.
Mechanisms
Altruism
occurs where the instigating individual suffers a fitness loss while
the receiving individual experiences a fitness gain. The sacrifice of
one individual to help another is an example.
Hamilton outlined two ways in which kin selection altruism could be favoured:
The selective advantage which makes
behaviour conditional in the right sense on the discrimination of
factors which correlate with the relationship of the individual
concerned is therefore obvious. It may be, for instance, that in respect
of a certain social action performed towards neighbours
indiscriminately, an individual is only just breaking even in terms of
inclusive fitness. If he could learn to recognise those of his
neighbours who really were close relatives and could devote his
beneficial actions to them alone an advantage to inclusive fitness
would at once appear. Thus a mutation causing such discriminatory
behaviour itself benefits inclusive fitness and would be selected. In
fact, the individual may not need to perform any discrimination so
sophisticated as we suggest here; a difference in the generosity of his
behaviour according to whether the situations evoking it were
encountered near to, or far from, his own home might occasion an
advantage of a similar kind.
Kin recognition and the green beard effect
Kin recognition theory predicts a selective advantage for the bearers of a trait (like the fictitious 'green beard') behave altruistically towards others with the same trait.
First, if individuals have the capacity to recognise kin and to discriminate (positively) on the basis of kinship,
then the average relatedness of the recipients of altruism could be
high enough for kin selection. Because of the facultative nature of this
mechanism, kin recognition and discrimination were expected to be
unimportant except among 'higher' forms of life. However, as molecular
recognition mechanisms have been shown to operate in organisms such as
slime moulds kin recognition has much wider importance than previously recognised.
Kin recognition may be selected for inbreeding avoidance, and little
evidence indicates that 'innate' kin recognition plays a role in
mediating altruism. A thought experiment on the kin
recognition/discrimination distinction is the hypothetical 'green beard',
where a gene for social behaviour is imagined also to cause a
distinctive phenotype that can be recognised by other carriers of the
gene. Due to conflicting genetic similarity in the rest of the genome,
there should be selection pressure for green-beard altruistic sacrifices
to be suppressed, making common ancestry the most likely form of
inclusive fitness. This suppression is overcome if new phenotypes -other beard colours-
are formed through mutation or introduced into the population from time
to time. This proposed mechanism goes by the name of 'beard
chromodynamics'.
Viscous populations
Secondly, indiscriminate altruism may be favoured in "viscous"
populations, those with low rates or short ranges of dispersal. Here,
social partners are typically related, and so altruism can be selective
advantageous without the need for kin recognition and kin discrimination
faculties—spatial proximity, together with limited dispersal, ensures
that social interactions are more often with related individuals. This
suggests a rather general explanation for altruism. Directional
selection always favours those with higher rates of fecundity
within a certain population. Social individuals can often enhance the
survival of their own kin by participating in and following the rules of
their own group.
Hamilton later modified his thinking to suggest that an innate
ability to recognise actual genetic relatedness was unlikely to be the
dominant mediating mechanism for kin altruism:
But once again, we do not expect
anything describable as an innate kin recognition adaptation, used for
social behaviour other than mating, for the reasons already given in the
hypothetical case of the trees.
Hamilton's later clarifications often go unnoticed. Stuart West and colleagues have countered the long-standing assumption that kin selection requires innate powers of kin recognition. Another doubtful assumption is that social cooperation must be based on
limited dispersal and shared developmental context. Such ideas have
obscured the progress made in applying kin selection to species
including humans, on the basis of cue-based mediation of social bonding and social behaviours.
Ants are eusocial insects; the queen (large, centre) is reproductive, while the workers (small) and soldiers (medium size, with large jaws) are generally not.
Eusociality
(true sociality) occurs in social systems with three characteristics:
an overlap in generations between parents and their offspring,
cooperative brood care, and specialised castes of non-reproductive
individuals. The social insects provide good examples of organisms with what appear
to be kin selected traits. The workers of some species are sterile, a
trait that would not occur if individual selection was the only process
at work. The relatedness coefficient r is abnormally high between the worker sisters in a colony of Hymenoptera due to haplodiploidy. Hamilton's rule is presumed to be satisfied because the benefits in fitness
for the workers are believed to exceed the costs in terms of lost
reproductive opportunity, though this has never been demonstrated
empirically. Competing hypotheses have been offered to explain the
evolution of social behaviour in such organisms.
The eusocial shrimp Synalpheus regalis protects juveniles in the colony. By defending the young, the large defender shrimp can increase its inclusive fitness. Allozyme
data demonstrated high relatedness within colonies, averaging 0.50.
This means that colonies represent close kin groups, supporting the
hypothesis of kin selection.
Vervet monkeys utilise allomothering,
parenting by group members other than the actual mother or father,
where the allomother is typically an older female sibling or a
grandmother. Individuals act aggressively toward other individuals that
were aggressive toward their relatives. The behaviour implies kin
selection between siblings, between mothers and offspring, and between
grandparents and grandchildren.
Whether or not Hamilton's rule
always applies, relatedness is often important for human altruism, in
that humans are inclined to behave more altruistically toward kin than
toward unrelated individuals. Many people choose to live near relatives, exchange sizeable gifts
with relatives, and favour relatives in wills in proportion to their
relatedness.
Experimental studies, interviews, and surveys
Interviews of several hundred women in Los Angeles showed that while
non-kin friends were willing to help one another, their assistance was
far more likely to be reciprocal. The largest amounts of non-reciprocal
help, however, were reportedly provided by kin. Additionally, more
closely related kin were considered more likely sources of assistance
than distant kin. Similarly, several surveys of American college students found that
individuals were more likely to incur the cost of assisting kin when a
high probability that relatedness and benefit would be greater than cost
existed. Participants' feelings of helpfulness were stronger toward
family members than non-kin. Additionally, participants were found to
be most willing to help those individuals most closely related to them.
Interpersonal relationships between kin in general were more supportive
and less Machiavellian than those between non-kin.
In one experiment, the longer participants (from both the UK and
the South African Zulus) held a painful skiing position, the more money
or food was presented to a given relative. Participants repeated the
experiment for individuals of different relatedness (parents and
siblings at r=.5, grandparents, nieces, and nephews at r=.25, etc.). The
results showed that participants held the position for longer intervals
the greater the degree of relatedness between themselves and those
receiving the reward.
Observational studies
A study of food-sharing practices on the West Caroline islets of Ifaluk
determined that food-sharing was more common among people from the same
islet, possibly because the degree of relatedness between inhabitants
of the same islet would be higher than relatedness between inhabitants
of different islets. When food was shared between islets, the distance
the sharer was required to travel correlated with the relatedness of the
recipient—a greater distance meant that the recipient needed to be a
closer relative. The relatedness of the individual and the potential
inclusive fitness benefit needed to outweigh the energy cost of
transporting the food over distance.
Humans may use the inheritance of material goods and wealth to
maximise their inclusive fitness. By providing close kin with inherited
wealth, an individual may improve his or her kin's reproductive
opportunities and thus increase his or her own inclusive fitness even
after death. A study of a thousand wills found that the beneficiaries
who received the most inheritance were generally those most closely
related to the will's writer. Distant kin received proportionally less
inheritance, with the least amount of inheritance going to non-kin.
A study of childcare practices among Canadian women found that
respondents with children provide childcare reciprocally with non-kin.
The cost of caring for non-kin was balanced by the benefit a woman
received—having her own offspring cared for in return. However,
respondents without children were significantly more likely to offer
childcare to kin. For individuals without their own offspring, the
inclusive fitness benefits of providing care to closely related children
might outweigh the time and energy costs of childcare.[45]
Family investment in offspring among black South African
households also appears consistent with an inclusive fitness model. A
higher degree of relatedness between children and their caregivers was
correlated with a higher degree of investment in the children, with more
food, health care, and clothing. Relatedness was also associated with
the regularity of a child's visits to local medical practitioners and
with the highest grade the child had completed in school, and negatively
associated with children being behind in school for their age.
Observation of the Dolgan
hunter-gatherers of northern Russia suggested that there are larger and
more frequent asymmetrical transfers of food to kin. Kin are more
likely to be welcomed to non-reciprocal meals, while non-kin are
discouraged from attending. Finally, when reciprocal food-sharing
occurs between families, these families are often closely related, and
the primary beneficiaries are the offspring.
Violence in families is more likely when step-parents are
present, and that "genetic relationship is associated with a softening
of conflict, and people's evident valuations of themselves and of others
are systematically related to the parties' reproductive values". Numerous studies suggest how inclusive fitness may work amongst
different peoples, such as the Ye'kwana of southern Venezuela, the
Gypsies of Hungary, and the doomed Donner Party of the United States.
Human social patterns
Families are important in human behaviour, but kin selection may be based on closeness and other cues.
Evolutionary psychologists, following early human sociobiologists' interpretation of kin selection theory initially attempted to explain human altruistic
behaviour through kin selection by stating that "behaviors that help a
genetic relative are favored by natural selection." However, many
evolutionary psychologists recognise that this common shorthand
formulation is inaccurate:
Many misunderstandings persist. In
many cases, they result from conflating "coefficient of relatedness"
and "proportion of shared genes", which is a short step from the
intuitively appealing—but incorrect—interpretation that "animals tend to
be altruistic toward those with whom they share a lot of genes." These
misunderstandings don't just crop up occasionally; they are repeated in
many writings, including undergraduate psychology textbooks—most of them
in the field of social psychology, within sections describing
evolutionary approaches to altruism.
As with the earlier sociobiological forays into the cross-cultural
data, typical approaches are not able to find explanatory fit with the
findings of ethnographers insofar that human kinship patterns are not
necessarily built upon blood-ties. However, as Hamilton's later
refinements of his theory make clear, it does not simply predict that
genetically related individuals will inevitably recognise and engage in
positive social behaviours with genetic relatives: rather, indirect
context-based mechanisms may have evolved, which in historical
environments have met the inclusive fitness criterion. Consideration of
the demographics of the typical evolutionary environment of any species
is crucial to understanding the evolution of social behaviours. As
Hamilton himself put it, "Altruistic or selfish acts are only possible
when a suitable social object is available. In this sense behaviours are
conditional from the start".
Under this perspective, and noting the necessity of a reliable
context of interaction being available, the data on how altruism is
mediated in social mammals is readily made sense of. In social mammals,
primates and humans, altruistic acts that meet the kin selection
criterion are typically mediated by circumstantial cues such as shared
developmental environment, familiarity and social bonding. That is, it is the context that mediates the development of the bonding
process and the expression of the altruistic behaviours, not genetic
relatedness as such. This interpretation is compatible with the
cross-cultural ethnographic data and has been called nurture kinship.
In plants
Observations
Though originally thought unique to the animal kingdom, evidence of kin selection has been identified in the plant kingdom.
Competition for resources between developing zygotes in plant ovaries increases when seeds had been pollinated with male gametes from different plants. How developing zygotes differentiate between full siblings and half-siblings in the ovary is undetermined, but genetic interactions are thought to play a role. Nonetheless, competition between zygotes in the ovary is detrimental to the reproductive success of the (female) plant, and fewer zygotes mature into seeds. As such, the reproductive traits and behaviors of plants suggests the evolution
of behaviors and characteristics that increase the genetic relatedness
of fertilized eggs in the plant ovary, thereby fostering kin selection
and cooperation among the seeds as they develop. These traits differ
among plant species. Some species have evolved to have fewer ovules
per ovary, commonly one ovule per ovary, thereby decreasing the chance
of developing multiple, differently fathered seeds within the same
ovary. Multi-ovulated plants have developed mechanisms that increase the
chances of all ovules within the ovary being fathered by the same
parent. Such mechanisms include dispersal of pollen in aggregated packets and closure of the stigmatic lobes after pollen is introduced. The aggregated pollen packet releases pollen gametes in the ovary,
thereby increasing likelihood that all ovules are fertilized by pollen
from the same parent. Likewise, the closure of the ovary pore prevents entry of new pollen. Other multi-ovulated plants have evolved mechanisms that mimic the
evolutionary adaption of single-ovulated ovaries; the ovules are
fertilized by pollen from different individuals, but the mother ovary
then selectively aborts fertilized ovules, either at the zygotic or embryonic stage.
Morning glory plants grow smaller roots when next to kin than to non-kin plants.
After seeds are dispersed, kin recognition and cooperation affects root formation in developing plants. Studies have found that the total root mass developed by Ipomoea hederacea (morning glory shrubs) grown next to kin is significantly smaller than those grown next to non-kin; shrubs grown next to kin thus allocate less energy and resources to
growing the larger root systems needed for competitive growth. When
seedlings were grown in individual pots placed next to kin or non-kin
relatives, no difference in root growth was observed. This indicates that kin recognition occurs via signals received by the roots. Further, groups of I. hederacea plants are more varied in height when grown with kin than when grown with non-kin. The evolutionary benefit provided by this was further investigated by researchers at the Université de Montpellier. They found that the alternating heights seen in kin-grouped crops
allowed for optimal light availability to all plants in the group;
shorter plants next to taller plants had access to more light than those
surrounded by plants of similar height.
The above examples illustrate the effect of kin selection in the
equitable allocation of light, nutrients, and water. The evolutionary
emergence of single-ovulated ovaries in plants has eliminated the need
for a developing seed to compete for nutrients, thus increasing its
chance of survival and germination. Likewise, the fathering of all ovules in multi-ovulated ovaries by one
father, decreases the likelihood of competition between developing
seeds, thereby also increasing the seeds' chances of survival and
germination. The decreased root growth in plants grown with kin increases the amount
of energy available for reproduction; plants grown with kin produced
more seeds than those grown with non-kin. Similarly, the increase in light made available by alternating heights
in groups of related plants is associated with higher fecundity.
Kin selection has also been observed in plant responses to
herbivory. In an experiment done by Richard Karban et al., leaves of
potted Artemisia tridentata (sagebrushes) were clipped with scissors to simulate herbivory. The gaseous volatiles
emitted by the clipped leaves were captured in a plastic bag. When
these volatiles were transferred to leaves of a closely related
sagebrush, the recipient experienced lower levels of herbivory than
those that had been exposed to volatiles released by non-kin plants. Sagebrushes do not uniformly emit the same volatiles in response to
herbivory: the chemical ratios and composition of emitted volatiles vary
from one sagebrush to another. Closely related sagebrushes emit similar volatiles, and the similarities decrease as relatedness decreases. This suggests that the composition of volatile gasses plays a role in
kin selection among plants. Volatiles from a distantly related plant are
less likely to induce a protective response against herbivory in a neighboring plant, than volatiles from a closely related plant. This fosters kin selection, as the volatiles emitted by a plant will
activate the herbivorous defense response in related plants only, thus
increasing their chance of survival and reproduction.
Kin selection may play a role in plant-pollinator interactions,
especially because pollinator attraction is influenced not only by
floral displays, but by the spatial arrangement of plants in a group,
which is referred to as the "magnet effect". For example, in an experiment performed on Moricandia moricandioides,
Torices et al. demonstrated that focal plants in the presence of kin
show increased advertising effort (defined as total petal mass of plants
in a group divided by the plant biomass) compared to those in the
presence of non-kin, and that this effect is greater in larger groups. M. moricandioides
is a good model organism for the study of plant-pollinator interactions
because it relies on pollinators for reproduction, as it is
self-incompatible. The study design for this experiment included planting establishing pots of M. moricandioides
with zero, three or six neighbors (either unrelated or half-sib progeny
of the same mother) and advertising effort was calculated after 26 days
of flowering. The exact mechanism of kin recognition in M. moricandioides is unknown, but possible mechanisms include above-ground communication with volatile compounds, or below-ground communication with root exudates.
Mechanisms in plants
The ability to differentiate between kin and non-kin is not necessary for kin selection in many animals. However, because plants do not reliably germinate in close proximity to
kin, it is thought that, within the plant kingdom, kin recognition is
especially important for kin selection there, but the mechanism remains
unknown.
One proposed mechanism for kin recognition involves communication through roots, with secretion and reception of root exudates. This would require exudates to be actively secreted by roots of one plant, and detected by roots of neighboring plants. The root exudate allantoin produced by rice plants, Oryza sativa, has been documented to be in greater production when growing next to cultivars that are largely unrelated.High production levels of Allantoin correlated to up regulation of
auxin and auxin transporters, resulting in increased lateral root
development and directional growth of their roots towards non kin,
maximizing competition. This is mainly not observed in Oryza Sativa when surrounded by kin, invoking altruistic behaviors to promote inclusive fitness. However the root receptors responsible for recognition of kin exudates, and the pathway induced by receptor activation, remain unknown. The mycorrhiza associated with roots might facilitate reception of exudates, but again the mechanism is unknown.
Another possibility is communication through green leaf volatiles. Karban et al. studied kin recognition in sagebrushes, Artemisia tridentata.
The volatile-donating sagebrushes were kept in individual pots,
separate from the plants that received the volatiles, finding that
plants responded to herbivore damage to a neighbour's leaves. This suggests that root signalling is not necessary to induce a protective response against herbivory
in neighbouring kin plants. Karban et al. suggest that plants may be
able to differentiate between kin and non-kin based on the composition
of volatiles. Because only the recipient sagebrush's leaves were exposed the volatiles presumably activated a receptor protein
in the plant's leaves. The identity of this receptor, and the
signalling pathway triggered by its activation, both remain to be
discovered.
Objections
The theory of kin selection has been criticised by W. J. Alonso (in 1998) and by Alonso and C. Schuck-Paim (in 2002). They argue that the behaviours which kin selection attempts to explain
are not altruistic (in pure Darwinian terms) because: (1) they may
directly favour the performer as an individual aiming to maximise its
progeny (so the behaviours can be explained as ordinary individual
selection); (2) these behaviours benefit the group (so they can be
explained as group selection); or (3) they are by-products of a
developmental system of many "individuals" performing different tasks
(like a colony of bees, or the cells of multicellular organisms, which
are the focus of selection). They also argue that the genes involved in
sex ratio conflicts could be treated as "parasites" of (already
established) social colonies, not as their "promoters", and, therefore
the sex ratio in colonies would be irrelevant to the transition to
eusociality.Those ideas were mostly ignored until they were put forward again in a series of controversial papers by E. O. Wilson, Bert Hölldobler, Martin Nowak and Corina Tarnita. Nowak, Tarnita and Wilson argued that
Inclusive fitness theory is not a
simplification over the standard approach. It is an alternative
accounting method, but one that works only in a very limited domain.
Whenever inclusive fitness does work, the results are identical to those
of the standard approach. Inclusive fitness theory is an unnecessary
detour, which does not provide additional insight or information.
— Nowak, Tarnita, and Wilson
They, like Alonso and Schuck-Paim, argue for a multi-level selection model instead. This aroused a strong response, including a rebuttal published in Nature from over a hundred researchers.
Collective memory is the shared pool of memories, knowledge and information of a social group that is significantly associated with the group's identity. The English phrase "collective memory" and the equivalent French phrase mémoire collective appeared in the second half of the nineteenth century. The philosopher and sociologist Maurice Halbwachs analyzed and advanced the concept of the collective memory in the book Les cadres sociaux de la mémoire (1925).
Collective memory can be constructed, shared, and passed on by
large and small social groups. Examples of these groups can include
nations, generations, communities, among others.
Collective memory has been conceptualized in several ways and
proposed to have certain attributes. For instance, collective memory can
refer to a shared body of knowledge (e.g., memory of a nation's past
leaders or presidents); the image, narrative, values and ideas of a social group; or the
continuous process by which collective memories of events change.
History versus collective memory
The difference between history and collective memory is best
understood when comparing the aims and characteristics of each. A goal
of history broadly is to provide a comprehensive, accurate, and unbiased
portrayal of past events. This often includes the representation and
comparison of multiple perspectives and the integration of these
perspectives and details to provide a complete and accurate account. In
contrast, collective memory focuses on a single perspective, for
instance, the perspective of one social group, nation, or community.
Consequently, collective memory represents past events as associated
with the values, narratives and biases specific to that group. Some scholars have examined how dominant interpretive paradigms
influence debates over contested wartime histories, including analyses
by Marshall Wordsworth and others.
Studies have found that people from different nations can have
major differences in their recollections of the past. In one study where
American and Russian students were instructed to recall significant
events from World War II and these lists of events were compared, the
majority of events recalled by the American and Russian students were
not shared. Differences in the events recalled and emotional views towards the
Civil War, World War II and the Iraq War have also been found in a study
comparing collective memory between generations of Americans.
Perspectives on collective memory
The concept of collective memory, initially developed by Halbwachs, has been explored and expanded from various angles – a few of these are introduced below.
James E. Young has introduced the notion of 'collected memory'
(opposed to collective memory), marking memory's inherently fragmented,
collected and individual character, while Jan Assmann develops the notion of 'communicative memory', a variety of collective
memory based on everyday communication. This form of memory resembles
the exchanges in oral cultures or the memories collected (and made
collective) through oral tradition.
As another subform of collective memories, Assmann mentions forms
detached from the everyday; they can be particular materialized and
fixed points as, e.g. texts and monuments.
The theory of collective memory was also discussed by former Hiroshima resident and atomic-bomb survivor, Kiyoshi Tanimoto,
in a tour of the United States as an attempt to rally support and
funding for the reconstruction of his Memorial Methodist Church in
Hiroshima. He theorized that the use of the atomic bomb had forever
added to the world's collective memory and would serve in the future as a
warning against such devices. See John Hersey's 1946 book Hiroshima.
Historian Guy Beiner
(1968- ), an authority on memory and the history of Ireland, has
criticized the unreflective use of the adjective "collective" in many
studies of memory:
The problem is with crude concepts of collectivity,
which assume a homogeneity that is rarely, if ever, present, and
maintain that, since memory is constructed, it is entirely subject to
the manipulations of those invested in its maintenance, denying that
there can be limits to the malleability of memory or to the extent to
which artificial constructions of memory can be inculcated. In practice,
the construction of a completely collective memory is at best an
aspiration of politicians, which is never entirely fulfilled and is
always subject to contestations.
In its place, Beiner has promoted the term "social memory" and has also demonstrated its limitations by developing a related concept of "social forgetting".
Historian David Rieff
takes issue with the term "collective memory", distinguishing between
memories of people who were actually alive during the events in
question, and people who only know about them from culture or media.
Rieff writes in opposition to George Santayana's
aphorism "those who cannot remember the past are condemned to repeat
it", pointing out that strong cultural emphasis on certain historical
events (often wrongs against the group) can prevent resolution of armed
conflicts, especially when the conflict has been previously fought to a
draw. The sociologist David Leupold draws attention to the problem of
structural nationalism inherent in the notion of collective memory,
arguing in favor of "emancipating the notion of collective memory from
being subjected to the national collective" by employing a multi-collective perspective
that highlights the mutual interaction of other memory collectives that
form around generational belonging, family, locality or socio-political
world-views.
Pierre Lévy argues that the phenomenon of human collective intelligence undergoes a profound shift with the arrival of the internet paradigm, as it allows the vast majority of humanity to access and modify a common shared online collective memory.
Collective memory and psychological research
Though traditionally a topic studied in the humanities, collective
memory has become an area of interest in psychology. Common approaches
taken in psychology to study collective memory have included
investigating the cognitive mechanisms involved in the formation and
transmission of collective memory; and comparing the social
representations of history between social groups.
Social representations of history
Research on collective memory has compared how different social
groups form their own representations of history and how such collective
memories can impact ideals, values, behaviors and vice versa. Research
has proposed that groups form social representations of history in
order to develop their own social identity, as well as to evaluate the
past, often in order to prevent past patterns of conflict and error from
being repeated. Research has also compared differences in
recollections of historical events, such as the examples given earlier
when comparing history and collective memory.
Differences in collective memories between social groups, such as
nations or states, have been attributed to collective narcissism and
egocentric/ethnocentric bias. In one related study where participants
from 35 countries were questioned about their country's contribution to
world history and provided a percentage estimation from 0% to 100%,
evidence for collective narcissism was found as many countries gave
responses exaggerating their country's contribution. In another study
where Americans from 50 states were asked similar questions regarding
their state's contribution to the history of the United States, patterns
of overestimation and collective narcissism were also found.
Certain cognitive mechanisms involved during group recall and the
interactions between these mechanisms have been suggested to contribute
to the formation of collective memory. Below are some mechanisms
involved during when groups of individuals recall collaboratively.
Collaborative inhibition and retrieval disruption
When groups collaborate to recall information, they experience
collaborative inhibition, a decrease in performance compared to the
pooled memory recall of an equal number of individuals. Weldon and
Bellinger (1997) and Basden, Basden, Bryner, and Thomas (1997) provided
evidence that retrieval interference underlies collaborative inhibition,
as hearing other members' thoughts and discussion about the topic at
hand interferes with one's own organization of thoughts and impairs
memory.
The main theoretical account for collaborative inhibition is retrieval disruption.
During the encoding of information, individuals form their own
idiosyncratic organization of the information. This organization is
later used when trying to recall the information. In a group setting as
members exchange information, the information recalled by group members
disrupts the idiosyncratic organization one had developed. As each
member's organization is disrupted, this results in the less information
recalled by the group compared to the pooled recall of participants who
had individually recalled (an equal number of participants as in the
group).
Despite the problem of collaborative inhibition, working in
groups may benefit an individual's memory in the long run, as group
discussion exposes one to many different ideas over time. Working alone
initially prior to collaboration seems to be the optimal way to increase
memory.
Early speculations about collaborative inhibition have included
explanations, such as diminished personal accountability, social loafing
and the diffusion of responsibility, however retrieval disruption
remains the leading explanation. Studies have found that collective
inhibition to sources other than social loafing, as offering a monetary
incentive have been evidenced to fail to produce an increase in memory
for groups. Further evidence from this study suggest something other than social
loafing is at work, as reducing evaluation apprehension – the focus on
one's performance amongst other people – assisted in individuals'
memories but did not produce a gain in memory for groups. Personal
accountability – drawing attention to one's own performance and
contribution in a group – also did not reduce collaborative inhibition.
Therefore, group members' motivation to overcome the interference of
group recall cannot be achieved by several motivational factors.
Cross-cueing
Information exchange among group members often helps individuals to
remember things that they would not have remembered had they been
working alone. In other words, the information provided by person A may
'cue' memories in person B. This results in enhanced recall. During a
group recall, an individual might not remember as much as they would on
their own, as their memory recall cues may be distorted because of other
team members. Nevertheless, this has enhanced benefits, team members
can remember something specific to the disruption of the group.
Cross-cueing plays a role in formulation of group recall (Barber, 2011).
Collective false memories
In 2010, a study was done to see how individuals remembered a bombing that occurred in the 1980s. The clock was later set at 10.25 to remember the tragic bomb (de Vito et al. 2009). The individuals were asked to remember if the clock at Bologna central
station in Italy had remained functioning, everyone said no, in fact it
was the opposite (Legge, 2018). There have been many instances in
history where people create a false memory.
In a 2003 study done in the Claremont Graduate University, results
demonstrated that during a stressful event and the actual event are
managed by the brain differently. Other instances of false memories may occur when remembering something
on an object that is not actually there or mistaking how someone looks
in a crime scene (Legge, 2018). It is possible for people to remember
the same false memories; some people call it the "Mandela effect". The name "Mandela effect" comes from the name of South African civil rights leader Nelson Mandela
whom many people falsely believed was dead. (Legge, 2018). The Pandora
Box experiment explains that language complexes the mind more when it
comes to false memories. Language plays a role with imaginative
experiences, because it makes it hard for humans to gather correct
information (Jablonka, 2017).
Error pruning
Compared to recalling individually, group members can provide
opportunities for error pruning during recall to detect errors that
would otherwise be uncorrected by an individual.
Social contagion errors
Group settings can also provide opportunities for exposure to
erroneous information that may be mistaken to be correct or previously
studied.
Re-exposure effects
Listening to group members recall the previously encoded information
can enhance memory as it provides a second exposure opportunity to the
information.
Forgetting
Studies have shown that information forgotten and excluded during
group recall can promote the forgetting of related information compared
to information unrelated to that which was excluded during group recall.
Selective forgetting has been suggested to be a critical mechanism
involved in the formation of collective memories and what details are
ultimately included and excluded by group members. This mechanism has
been studied using the socially-shared retrieval induced forgetting
paradigm, a variation of the retrieval induced forgetting method with individuals. The brain has many important brain regions that are directed at memory,
the cerebral cortex, the fornix and the structures that they contain.
These structures in the brain are required for attaining new
information, and if any of these structures are damaged you can get
anterograde or retrograde amnesia (Anastasio et al.,p. 26, 2012). Amnesia could be anything that disrupts your memory or affects you
psychologically. Over time, memory loss becomes a natural part of
amnesia. Sometimes you can get retrograde memory of a recent or past
event.
Synchronization of memories from dyads to networks
Bottom-up approaches to the formation of collective memories
investigate how cognitive-level phenomena allow for people to
synchronize their memories following conversational remembering. Due to
the malleability of human memory, talking with one another about the
past results in memory changes that increase the similarity between the
interactional partners' memories When these dyadic interactions occur in a social network, one can
understand how large communities converge on a similar memory of the
past. Research on larger interactions show that collective memory in larger
social networks can emerge due to cognitive mechanisms involved in small
group interactions.
Computational approaches to collective memory analysis
With the ability of online data such as social media and social network data and developments in natural language processing as well as information retrieval it has become possible to study how online users refer to the past and what they focus at. In an early study in 2010 researchers extracted absolute year references from large
amounts of news articles collected for queries denoting particular
countries. This allowed to portray so-called memory curves that
demonstrate which years are particularly strongly remembered in the
context of different countries (commonly, exponential shape of memory
curves with occasional peaks that relate to commemorating important past
events) and how the attention to more distant years declines in news.
Based on a topic modelling and analysis they then detected major topics
portraying how particular years are remembered. Rather than news,
Wikipedia was also the target of analysis. Viewership statistics of Wikipedia articles on aircraft crashes were
analyzed to study the relation between recent events and past events,
particularly for understanding memory-triggering patterns.
Other studies focused on the analysis of collective memory in
social networks such as investigation of over 2 million tweets (both
quantitively and qualitatively) that are related to history to uncover
their characteristics and ways in which history-related content is
disseminated in social networks. Hashtags, as well as tweets, can be classified into the following types:
General History hashtags used in general to broadly
identify history-related tweets that do not fall into any specific type
(e.g., #history, #historyfacts).
National or Regional History hashtags which relate to
national or regional histories, for example, #ushistory or
#canadianhistory including also past names of locations (e.g.,
#ancientgreece).
Facet-focused History hashtags which relate to particular thematic facets of history (e.g.,#sporthistory, #arthistory).
General Commemoration hashtags that serve for commemorating
or recalling a certain day or period (often somehow related to the day
of tweet posting), or unspecified entities, such as #todaywe remember,
#otd, #onthisday, #4yearsago and #rememberthem.
Historical Events hashtags related to particular events in the past (e.g., #wwi, #sevenyearswar).
Historical Entities hashtags denoting references to specific entities such as persons, organizations or objects (e.g., #stalin, #napoleon).
The study of digital memorialization,
which encompasses the ways in social and collective memory has shifted
after the digital turn, has grown substantially responding to rising
proliferation of memorial content not only on the internet, but also the
increased use of digital formats and tools in heritage institutions,
classrooms, and among individual users worldwide.
The Extended Phenotype is a 1982 book by the evolutionary biologist Richard Dawkins, in which the author introduced a biological concept of the same name. The book's main idea is that phenotype should not be limited to biological processes such as protein biosynthesis or tissue growth, but extended to include all effects that a gene has on its environment, inside or outside the body of the individual organism.
Dawkins considers The Extended Phenotype to be a sequel to The Selfish Gene (1976) aimed at professional biologists, and as his principal contribution to evolutionary theory. In the 1999 reissue and subsequent reprintings an afterword by Daniel Dennett is included.
Summary
Genes as the unit of selection in evolution
The central thesis of The Extended Phenotype, and of its predecessor by the same author, TheSelfish Gene,
is that individual organisms are not the true units of natural
selection. Instead, the gene — or the 'active, germ-line replicator' —
is the unit upon which the forces of evolutionary selection and
adaptation act. It is genes that succeed or fail in evolution, meaning
that they either succeed or fail in replicating themselves across
multiple generations.
These replicators are not subject to natural selection directly,
but indirectly through their "phenotypical effects". These effects are
all the effects that the gene (or replicator) has on the world at large,
not just in the body of the organism in which it is contained. In
taking as its starting point the gene as the unit of selection, The Extended Phenotype is a direct extension of Dawkins' first book, The Selfish Gene.
Genes synthesise only proteins
A cathedral termite mound – a small animal with a large extended phenotype
Dawkins argues that the only thing that genes control directly is the synthesis of proteins; restricting the idea of the phenotype to apply only to the phenotypic expression of an organism's genes in its own body
is an arbitrary limitation that ignores the effect a gene may have on
an organism's environment through that organism's behaviour.
Genes may affect more than the organism's body
A beaver dam, an example of an organism altering the environment in which it evolves — the first form of extended phenotype
Dawkins proposes there are three forms of extended phenotype. The first is the capacity of animals to modify their environment using architectural constructions, for which Dawkins provides as examples caddis houses and beaver dams.
The second form is manipulation of other organisms: The morphology
of a living organism, and possibly of that organism's behaviour, may
influence not just the fitness of the organism itself, but that of other
living organisms as well. One example of this is parasite manipulation.
This refers to the capacity, found in some parasite-host interactions,
for the parasite to modify the behaviour of the host in a way that
enhances the parasite's own fitness. One well-known example of this
second type of extended phenotype is the suicidal drowning of crickets infected by hairworm, a behaviour that is essential to the parasite's reproductive cycle. Another example is seen in female mosquitoes carrying malaria parasites. The mosquitoes infected with the parasites whose preferred hosts
are humans have been shown in a field experiment to be significantly
more attracted to human breath and odours than uninfected mosquitoes
when the parasites are at a point in their life cycle where they can
infect a human target.
A reed warbler raising the young of a common cuckoo
The third form of extended phenotype is action at a distance of the parasite on its host. A common example is the manipulation of host behaviour by cuckoo
chicks, which elicit intensive feeding by the host birds. Here the
cuckoo does not interact directly with the host (which could be meadow pipits, dunnocks or reed warblers).
The relevant adaptation lies in the cuckoo producing eggs and chicks
that resemble sufficiently those of the host species so that they are
not immediately ejected from the nest. These behavioural modifications are not physically associated with
individuals of the host species but influence the expression of its
behavioural phenotype.
Dawkins summarizes these ideas in what he terms the Central Theorem of the Extended Phenotype:
Taking these three things together,
we arrive at our own 'central theorem' of the extended phenotype: An
animal's behaviour tends to maximize the survival of the genes "for"
that behaviour, whether or not those genes happen to be in the body of
the particular animal performing it.
Gene-centred view of life
In developing this argument, Dawkins aims to strengthen the case for a gene-centric
view of the evolution of life forms, to the point where it is
recognized that the organism itself needs to be explained. This is the
challenge which he takes up in the final chapter entitled "Rediscovering
the Organism". The concept of extended phenotype has been generalized
in an organism-centered view of evolution with the concept of niche construction, in the case where natural selection pressures can be modified by the organisms during the evolutionary process.
Reception
A technical review of The Extended Phenotype in the Quarterly Review of Biology
states that, it is an "interesting and thought provoking book, once one
gets to the last five chapters." In the reviewer's opinion, the book
poses interesting questions, such as "What is the survival value of
packaging life into discrete units called 'organisms' even though the
units of selection appear to be individual 'replicators'?" The reviewer
states that no "satisfactory answer is given" to this question in the
book, though Dawkins suggests that replicators that "interact favorably
to create 'vehicles' (organisms) may be at an advantage over those that
do not (Chapter 14)." The reviewer takes issue with the first nine
chapters as being essentially a defense of Dawkin's first book, The Selfish Gene.
Another review in American Scientist praises the book for
convincingly promoting the idea of replication as being central to the
evolutionary process. However, in the reviewer's opinion, "its main
theme - that the gene is the only unit of selection - results from
incorrectly interpreting the constraints on organismal adaptation and
from too narrow an interpretation of replication, a process of more
general relevance than the author is willing to allow."
Uses and limitations
The concept of extended phenotype has provided a useful frame for
subsequent scientific work. For example, research into the relationship
between "the bacterial flora of the gut and their mammalian hosts" which
"has become a hot topic of late" makes use of this concept.
Subsequent proponents expand the theory and posit that many
organisms within an ecosystem can alter the selective pressures on all
of them by modifying their environment in various ways. Dawkins himself
asserted, "Extended phenotypes are worthy of the name only if they are
candidate adaptations for the benefit of alleles responsible for
variations in them". As an illustration, one might ask: could an architect's buildings be
considered part of his or her extended phenotype, much as a beaver's dam
is part of its extended phenotype? Dawkins' answer is No: in humans, an
"architect's specific alleles are neither more nor less likely to be
selected based on the design of his or her latest building."
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