Ombudsmen also aim to identify systemic issues leading to poor
service or breaches of people's rights. At the national level, most
ombudsmen have a wide mandate to deal with the entire public sector, and
sometimes also elements of the private sector (for example, contracted
service providers). In some cases, there is a more restricted mandate to
a certain sector of society. More recent developments have included the
creation of specialized children's ombudsmen.
In some countries, an inspector general,
citizen advocate or other official may have duties similar to those of a
national ombudsman and may also be appointed by a legislature. Below
the national level, an ombudsman may be appointed by a state, local, or
municipal government. Private ombudsmen may be appointed by, or even
work for, a corporation such as a utility supplier, newspaper, NGO, or professional regulatory body.
In some jurisdictions, an ombudsman charged with handling
concerns about national government is more formally referred to as the
"parliamentary commissioner" (e.g. the United Kingdom Parliamentary Commissioner for Administration, and the Western Australian state Ombudsman). In many countries where the ombudsman's responsibility includes protecting human rights, the ombudsman is recognized as the national human rights institution.
The post of ombudsman had by the end of the 20th century been
instituted by most governments and by some intergovernmental
organizations such as the European Union.
As of 2005, including national and sub-national levels, a total of 129
offices of ombudsman have been established around the world.
Origins and etymology
A prototype of an ombudsman may have flourished in China during the Qin dynasty (221 BC), and later in Korea during the Joseon dynasty. The position of secret royal inspector, or amhaeng-eosa (암행어사, 暗行御史)
was unique to the Joseon dynasty, where an undercover official directly
appointed by the king was sent to local provinces to monitor government
officials and look after the populace while travelling incognito. The Roman tribune had some similar roles, with the power to veto acts that infringed upon the Plebeians. Another precursor to the ombudsman was the Diwān al-Maẓālim (دِيوَانُ الْمَظَالِمِ) which appears to go back to the second caliph, Umar (634–644), and the concept of Qaḍī al-Quḍāt (قَاضِي الْقُضَاةِ). They were also attested in Siam, India, the Liao dynasty, Japan, and China.
An indigenous Swedish, Norwegian, and Danish term, ombudsman, ombodsmann, ombudsmand or ombudsmann is etymologically rooted in the Old Norse word umboðsmaðr, essentially meaning 'representative' (with the word umbud/ombod/ombud
meaning 'proxy', 'attorney'; that is, someone who is authorized to act
for someone else, a meaning it still has in the Scandinavian languages).
From 1552, it is used in the Nordic languages as the Swedish ombudsman (pronounced[ˈɔ̂mːbʉːdsˌman]ⓘ), the Danish ombudsmand, the Icelandic and Faroeseumboðsmaður and the Norwegian ombudsmann/ombodsmann.
The general meaning was and is approximately 'a man representing
(someone)' (i.e., a representative) or 'a man with a commission (from
someone)' (a commissioner). The Swedish-speaking minority in Finland
uses the Swedish terminology. The various forms of the suffix -mand, -maður, et cetera, are just the forms the common Germanic word represented by the English word man have in the various languages. Thus, the modern plural form ombudsmen of the English borrowed word ombudsman is likely.
Use of the term in its modern sense began in Sweden with the Swedish Parliamentary Ombudsman instituted by the Instrument of Government of 1809, to safeguard the rights of citizens by establishing a supervisory agency independent of the executive branch. The predecessor of the Swedish Parliamentary Ombudsman was the Office of Supreme Ombudsman (Högste Ombudsmannen), which was established by the Swedish King, Charles XII, in 1713. Charles XII was in exile in Turkey
and needed a representative in Sweden to ensure that judges and civil
servants acted in accordance with the laws and with their duties. If
they did not do so, the Supreme Ombudsman had the right to prosecute
them for negligence. In 1719 the Swedish Office of Supreme Ombudsman
became the Chancellor of Justice. The Parliamentary Ombudsman was established in 1809 by the Swedish Riksdag, as a parallel institution to the still-present Chancellor of Justice, reflecting the concept of separation of powers as developed by Montesquieu.
The Parliamentary Ombudsman is the institution that the
Scandinavian countries subsequently developed into its contemporary
form, and which subsequently has been adopted in many other parts of the
world. The word ombudsman and its specific meaning have since been
adopted in various languages, such as Dutch. The German language uses Ombudsmann, Ombudsfrau and Ombudsleute. Notable exceptions are French, Italian, Spanish, and Finnish, which use translations instead. Modern variations of this term include ombud, ombuds, ombudsperson, or ombudswoman, and the conventional English plural is ombudsmen. In Nigeria, the ombudsman is known as the Public Complaints Commission or the ombudsman.
In politics
In general, an ombudsman is a state official appointed to provide a
check on government activity in the interests of the citizen and to
oversee the investigation of complaints of improper government activity
against the citizen. If the ombudsman finds a complaint to be
substantiated, the problem may get rectified, or an ombudsman report is
published making recommendations for change. Further redress depends on
the laws of the country concerned, but this typically involves financial
compensation. Ombudsmen in most countries do not have the power to
initiate legal proceedings or prosecution on the grounds of a complaint.
This role is sometimes referred to as a "tribunician" role, and has
been traditionally fulfilled by elected representatives – the term
refers to the ancient Roman "tribunes of the plebeians" (tribuni plebis), whose role was to intercede in the political process on behalf of common citizens.
The significant advantage of an ombudsman is that they examine
complaints from outside the offending state institution, thus avoiding
the conflicts of interest inherent in self-policing. However, the
ombudsman system relies heavily on the selection of an appropriate
individual for the office, and on the cooperation of at least some
effective official from within the apparatus of the state. However,
sociologist Jürgen Beyer has criticised the institution, stating:
"Ombudsmen are relics of absolutism, designed to iron out the worst
excesses of administrative arbitrariness while keeping the power
structures intact."
Many private companies, universities, non-profit organisations, and
government agencies also have an ombudsman (or an ombuds office) to
serve internal employees, managers and/or other constituencies. These
ombudsman roles are structured to function independently, by reporting
to the CEO or board of directors, and, according to the International
Ombudsman Association (IOA) Standards of Practice, they do not have any
other role in the organisation. Organisational ombudsmen often receive
more complaints than alternative procedures such as anonymous hot-lines.
Since the 1960s, the profession has grown in the United States,
and Canada, particularly in corporations, universities, and government
agencies. The organizational ombudsman
works as a designated neutral party, one who is high-ranking in an
organization, but who is not part of executive management. Using an alternative dispute resolution (ADR) or appropriate dispute resolution approach, an organisational ombudsman can provide options to whistleblowers or employees and managers with ethical
concerns; provide coaching, shuttle diplomacy, generic solutions
(meaning a solution which protects the identity of one individual by
applying to a class of people, rather than just for the one individual)
and mediation
for conflicts; track problem areas; and make recommendations for
changes to policies or procedures in support of orderly systems change.
Darwinian anthropology describes an approach to anthropological analysis which employs various theories from Darwinianevolutionary biology. Whilst there are a number of areas of research that can come under this broad description some specific research projects have been closely associated with the
label. A prominent example is the project that developed in the mid
1970s with the goal of applying sociobiological perspectives to explain patterns of human social relationships, particularly kinship patterns across human cultures.
This kinship-focused Darwinian anthropology was a significant intellectual forebear of evolutionary psychology, and both draw on biological theories of the evolution of social behavior (in particular inclusive fitness theory) upon which the field of sociobiology was founded.
Although ten years have passed
since Hamilton's landmark papers, apparently only a single social
scientist (Campbell, 31) has made a distinct effort to incorporate kin
selection into theories of human altruism... But so have the
biologists, for one reason or another, failed to consider the enormous
literature on topics like kinship systems and reciprocity in human
behavior.
Amongst other suggestions, Alexander suggested that certain patterns
of social cooperation documented by ethnographers, in particular the avunculate
('mother's brother') relationship, could be explained in reference to
individuals pursuing a strategy of individual inclusive fitness
maximization under conditions of low certainty-of-paternity. This
hypothesis was subsequently taken up and elaborated in a series of
studies by other Darwinian anthropologists:
The hypothesis follows that
matrilineal inheritance is a cultural trait that evolved in response to
low probability of paternity. [...] The paternity hypothesis was
rescued and made explicit in the context of modern evolutionary
theory by Alexander in 1974. Over the next 10 years this provided the
impetus leading to numerous theoretical refinements and scholarly
and empirical investigations (Flinn 1981; Gaulin & Schlegel
1980; Greene 1978; Hartung 1981b; Kurland 1979).(Hartung
1985,661-663)
Ultimately these analyses were considered unsuccessful, and were
specifically criticized by other sociobiologists on a number of grounds.
One problem was said to be that interpreting inclusive fitness
theory to imply that individuals have evolved the characteristic of
pursuing strategies to maximize their own 'inclusive fitness' is
erroneous; the theory should instead be interpreted to describe
selection pressures on genes:
Alexander's argument... erred through looking at things from the point of view of an individual...
I believe this kind of error is all too easy to make when we use the
technical term ‘fitness’ [of individuals]. This is why I have avoided
using the term in this book. There is really only one entity whose
point of view matters in evolution, and that entity is the selfish
gene.(Dawkins 1989 (1976), 137 emphasis in original)
A related problem was that, in assuming that individuals simply
operate as inclusive fitness maximizing agents, any investigation of the
proximate psychological mechanisms that mediate social behaviors was
ignored. Symons made this observation in his 1989 Critique of Darwinian Anthropology:
DA's central hypothesis is that
“evolved behavioral tendencies” cause human “behavior to assume
the form that maximizes inclusive fitness”(Irons 1979b, 33).
Turke and Betzig (1985) state this hypothesis as a formal
prediction: “Modern Darwinian theory predicts that human behavior will
be adaptive, that is, designed to promote maximum reproductive
success through available descendant and nondescendant relatives.”(p
79)… [T]he key terms in [this] quotation are used in DA to bypass the
question of phenotypic design in characterizations of
adaptation.(Symons 1989, 131-132)
Symons, along with Tooby and Cosmides, were amongst the founders of the emerging school of evolutionary psychology, whose aim has been to focus more on these hypothesized proximate psychological mechanisms.
Theoretical background
Darwinian anthropology was critiqued by Symonds for its agnosticism
as to the psychological mechanisms governing how social behavior is
actually expressed in the human species, and its reliance on
interpreting inclusive fitness theory to simply imply that humans have evolved to be inclusive fitness maximizers.
This section will review some of the relevant background discussion in
inclusive fitness theory to clarify why this position was considered
untenable.
Inclusive Fitness theory has often been interpreted to mean that social behavior per se is a goal of evolution,
and also that genes (or individual organisms) are selected to find
ways of actively distinguishing the identity of close genetic relatives
‘in order to’ engage in social behaviors with them.
[M]any 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. (Park 2007, p860)
The apparent rationale for this common mis-interpretation is that organisms would
thereby benefit the “Inclusive Fitness of the individuals (and
genes) involved”. This approach overlooks the point that evolution is
not a teleological process, but a passive, consequential and undirected biological process, where environmental variations and drift effects are present alongside random gene mutations and natural selection.
Inclusive fitness theory takes the form of an ultimate explanation, specifically a criterion (br>c), for the evolution of social behaviors, not a proximate mechanism governing the expression of social behaviors. What forms of social behavior might meet this criterion are cannot be a priori
specified by the theory, nor can it shed light on whether the life
history of a species provides opportunities for social interactions to
occur. Thus, strictly speaking, the interpretation that organisms ‘have
evolved to’ direct social behavior towards genetic relatives is not
implied by the theory (see also inclusive fitness).
Investigating how inclusive fitness theory might apply to the
potential emergence of social traits in any given species must begin
with an analysis of the evolutionarily typical ecological niche,
demographics, and patterns of interaction of that species. Where
significant interaction between individuals is not present in the life history of a species, the theory is necessarily null regarding social behaviors between individuals. As Silk (2001) put it;
The role of kinship in the daily
lives of animals depends on the demographic composition of the groups in
which they live. Kin selection will only be an important force in
the evolution of social behavior if animals find themselves in
situations where they have an opportunity to fulfill the
predictions of Hamilton's rule. At a minimum, kin must be
available. The number, availability, and degree of relatedness
among kin will depend on how groups are constructed in nature.”
(Silk 2001, 77)
Consideration must thus be given to whether the ecological niche
leads to the clustering of individuals in groups or whether individuals
are typically solitary. Socioecology
research, for example, suggests that fundamental influences on
demographic patterns are the distribution/fecundity of primary food
sources as well as patterns of predation. When considering social
behavior traits of a given species, consideration of these influences is
in a sense, logically prior to analyses of inclusive fitness pressures
on the species.
Selection pressure on genes or strategy of individuals
Darwinian anthropology, following R. D. Alexander, used the notion of the inclusive fitness of individuals rather than the inclusive fitness of genes.
Dawkins (above) pointed to this as an error. The source of the
confusion can be traced to discussions in Hamilton's early papers on
inclusive fitness. In his 1963 paper Hamilton refers, unambiguously, to
selection pressures on genes;
[T]he ultimate criterion which
determines whether G [a gene] will spread is not whether the
behaviour is to the benefit of the behaver but whether it is to the
benefit of the gene G; and this will be the case if the average net
result of the behaviour is to add to the gene pool a handful of
genes containing G in higher concentration than does the gene pool
itself.” (1996 [1963], 7)
However, in his paper published in 1964, actually written before the
1963 paper (Hamilton 1996), Hamilton had included a subsidiary
discussion on what the genetic theory might imply for how we look at the
fitness of individuals:
Actually, in the preceding
mathematical account we were not concerned with the inclusive
fitness of individuals as described here but rather with certain
averages of them which we call the inclusive fitness of types. But the
idea of the inclusive fitness of an individual is nevertheless a useful
one. Just as in the sense of classical selection we may consider whether
a given character expressed in an individual is adaptive in the sense
of being in the interest of his personal fitness or not, so in the
present sense of selection we may consider whether the character or
trait of behaviour is or is not adaptive in the sense if being in the
interest of his inclusive fitness.” (Hamilton 1996 [1964], 38)
It is clear here that the formal treatment is of the selection
pressures on types (genes or traits), whilst the notion of individual
inclusive fitness may serve as a guide to the adaptiveness of the trait;
just as consideration of effects of a trait on an individual's fitness
can be instructive when considering classical selection on traits. At
the same time, it is understandable that Alexander took the inclusive
fitness of individuals as a heuristic device.
Context-based or discrimination-based expression mechanisms
In his 1964 paper, Hamilton 'hazards' “the following unrigorous
statement of the main principle that has emerged from the model”;
The Social behaviour of a
species evolves in such a way that in each distinct
behaviour-evoking situation the individual will seem to value his
neighbours’ fitness against his own according to the coefficients
of relationship appropriate to that situation.”(1964 [1996], 49)
He uses the terms 'hazards', 'unrigorous', and 'will seem'
deliberately, since his formal analysis makes clear that the model
specifies the evolutionary selection pressure, rather than specifying
what mechanisms govern the proximate expression of social behaviors. He
also clearly points to social behaviors being evoked in distinct
situations, and that individuals may encounter potential social
recipients of different degree of relationship in different situations.
If one ignores the cautious qualifying words however, the passage might
readily be interpreted to imply that individuals are indeed expected to
make an active assessment of the degree of relatedness of others they
interact with in different situations. Later in the paper, Hamilton
again discusses the issue of whether the performance (or expression) of
social behaviors might be conditional on; (a) discriminating factors
which correlate with close relationship with the recipient, or (b)
actually discriminating which individuals 'really are' in close
relationship with the recipient:
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.” (1996 [1964], 51)
For certain social behaviors, Hamilton suggests there may be
selection pressure for more discerning discrimination of genetic
relatedness, were the mutation to occur. But 'in fact' the same net
result of accurately targeting social behaviors towards genetic
relatives could be achieved via a simpler mechanism of being expressed
in proximity to the actor's 'home'. Hamilton is thus agnostic as to
whether evolved social behaviors might be expressed via straightforward
proximate mechanisms such as location-based cues, or whether more
specific discriminatory powers might govern their expression. He does
suggest that the distinct social contexts within which various social
behaviors are expressed are factors to consider. Other theorists have
discussed these questions of whether proximity, context or more
discriminatory expression may govern behaviors:
Animals cannot, of course, be
expected to know, in a cognitive sense, who their relatives are, and in
practice the behaviour that is favoured by natural selection will be
equivalent to a rough rule of thumb such as ‘share food with anything
that moves in the nest in which you are sitting.’ If families happen to
go around in groups, this fact provides a useful rule of thumb for kin
selection: ‘care for any individual you often see’.” (Dawkins 1979, 187)
If, for example, animals behave with an equal degree of altruism to all their “neighbours”… and if on average
animals are related to their neighbours, then I would regard this as an
example of kin selection. It is not a necessary feature of kin
selection that an animal should distinguish different degrees of
relationship among its neighbours, and behave with greater altruism to
the more closely related…”(Maynard Smith 1976, 282 emphasis in original)
Dawkins believes social behaviors will in practice be governed by
context-based expression. Maynard Smith is, like Hamilton, agnostic, but
reiterates the point that context-based cues might well govern their
expression and that actively distinguishing relatives is not necessarily
expected for the expression of those social traits whose evolution is
governed by inclusive fitness criteria. In sum, inclusive fitness theory
does imply that; the evolutionary emergence of social behavior can occur where there is statistical association of genes between social actors and recipients; but that the expression
of such evolved social behaviors is not necessarily governed by actual
genetic relatedness between participants. The evolutionary criterion and
the proximate mechanism must thus not be confused: the first does
require genetic association (of the form br>c), the second does not.
Darwinian anthropology's central premise that human behavior has evolved to maximize the inclusive fitness of individuals is thus not a logical derivative of the theory. Also, the notion that humans will discriminate social behaviors towards genetic relatives is again not entailed by the theory.
Reception by anthropologists
Before the questions raised within anthropology about the study of ‘kinship’ by Schneider and others from the 1960s onwards, anthropology itself had paid very
little attention to the notion that social bonds were anything other
than connected to consanguinal (or genetic) relatedness (or its local
cultural conceptions). The social bonding associated with provision
of and sharing of food was one important exception, particularly in the
work of Richards, but this was largely ignored by descriptions of ‘kinship’ till more
recently. Although questioning the means by which ‘kinship bonds’ form,
few of these early accounts questioned the fundamental role of
‘procreative ties’ in social bonding (Schneider, 1984). From the 1950s
onwards, reports on kinship patterns in the New Guinea Highlands added
some momentum to what had until then been only occasional fleeting
suggestions that living together (co-residence) might underlie social
bonding, and eventually contributed to the general shift away from a
genealogical approach. For example, on the basis of his observations,
Barnes suggested:
[C]learly, genealogical connexion
of some sort is one criterion for membership of many social groups. But
it may not be the only criterion; birth, or residence, or a parent's
former residence, or utilization of garden land, or participation in
exchange and feasting activities or in house-building or raiding, may be
other relevant criteria for group membership.”(Barnes 1962,6)
Similarly, Langness' ethnography of the Bena Bena also emphasized a break with the genealogical perspective:
The sheer fact of residence in a Bena Bena group can and does determine kinship.
People do not necessarily reside where they do because they are
kinsmen: rather they become kinsmen because they reside there.”
(Langness 1964, 172 emphasis in original)
By 1972, Schneider had raised deep problems with the notion that human social bonds and
'kinship' was a natural category built upon genealogical ties (for more
information, see kinship), and especially in the wake of his 1984 critique this has become broadly accepted by most, if not all, anthropologists.
The darwinian anthropology (and other sociobiological)
perspectives, arising in the early 1970s, had not unreasonably assumed
that the genealogical conceptions of human kinship, in place since
Morgan's early work in the 1870s, were still valid as a universal feature of humans. But
they emerged at precisely the time that anthropology, being particularly
sensitive about its own apparent 'ethnocentric' generalizations about
kinship (from cultural particulars to human universals) was seeking to
distance itself from these conceptions. The vehemence of Sahlins'
rebuttal of sociobiology's genetic relatedness perspective in his 1976 The use and abuse of Biology, which underlined the non-genealogical nature of human kinship, can be understood as part of this 'distancing' trend.
Alternative approaches
The lack of success of darwinian anthropology created space for
alternative approaches to analyzing human social behaviors from a
biological perspective. Alexander's initial point (above) that the
inclusive fitness framework had been scarcely applied to human kinship
and social patterns has remained largely valid. But the move away from
genealogical kinship in anthropology has continued to be a major barrier
to any potential resolution. This section reviews a range of approaches
to synthesizing ideas from evolutionary biology to observations and
data about human social behaviour across contemporary human populations.
Whilst some of these approaches include the inclusive fitness approach,
others may seek to demonstrate fit to other theories from evolutionary
biology, or to demonstrate that certain proximate mechanisms of social
behaviour are both compatible with the inclusive fitness approach, and
also with the broad variety of ethnographic data on human kinship
patterns.
Theories in evolutionary biology relevant to understanding social
behavior may not be limited to frameworks such as inclusive fitness
theory. The theory of reciprocal altruism may have equal or greater explanatory power
for some forms of human social behavior, and perhaps kinship patterns.
Other approaches may maintain that human behavior is less amenable to
biological analysis due to the prominent influence of social learning
and cultural transmission in the human species, and instead advance
ideas based on the role of e.g. culture, historical contingencies or
economic/environmental conditions. All or any of these may or may not
contribute valuable insights to our understanding of social behavior and
social patterns in humans.
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.
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