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The Nestor effect: Extending evolutionary developmental psychology to a lifespan perspective

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Abstract

We extend an evolutionary perspective of development to the lifespan, proposing that human longevity may be related to the experience, knowledge, and wisdom provided by older members of human groups. In addition to the assistance in childcare provided by grandmothers to their daughters, the experience of wise elders could have served to benefit kin as well as members of the immediate group, and thus been the target of natural selection. We refer to the fitness-enhancing effect of cumulative and integrative knowledge of some members of a group as the Nestor effect, after the oldest member of the group of Greek kings in the siege of Troy. Furthermore, we argue that the contributions of such Nestors resulted in cultural innovations that in turn may have promoted care and respect for the elderly. This bidirectional effect makes culture both the product and producer of longevity.
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Review
The Nestor effect: Extending evolutionary developmental
psychology to a lifespan perspective
Werner Greve
*
, David F. Bjorklund
University of Hildesheim, Institute for Psychology, Marienburger Platz 22, D-31141 Hildesheim, Germany
article info
Article history:
Received 19 September 2008
Revised 1 April 2009
Available online 13 June 2009
Keywords:
Aging
Wisdom
Epigenetic inheritance
Adaptation
Cultural evolution
abstract
We extend an evolutionary perspective of development to the life-
span, proposing that human longevity may be related to the expe-
rience, knowledge, and wisdom provided by older members of
human groups. In addition to the assistance in childcare provided
by grandmothers to their daughters, the experience of wise elders
could have served to benefit kin as well as members of the imme-
diate group, and thus been the target of natural selection. We refer
to the fitness-enhancing effect of cumulative and integrative
knowledge of some members of a group as the Nestor effect, after
the oldest member of the group of Greek kings in the siege of Troy.
Furthermore, we argue that the contributions of such Nestors
resulted in cultural innovations that in turn may have promoted
care and respect for the elderly. This bidirectional effect makes cul-
ture both the product and producer of longevity.
Ó2009 Elsevier Inc. All rights reserved.
Some life experiences are, though potentially very useful, rarely used. Their value is high but hid-
den, as it were, like the back-up copy of your hard drive: you do not need it often, and it is likely that
you will never need it, but if you do need it, you need it badly. Cumulated experience, transmitted by
oral tradition is sometimes this type of life experience. As a recent example consider the Moken peo-
ple, so-called sea gypsies living on the islands off the coast of Thailand and Burma. Although this area
was devastated by the tsunami on December 26, 2004, they survived this catastrophe unscathed. The
Moken did not have expensive advanced technology to warn them about killer waves. They survived
merely because of their close relationships with and observation of nature and because they heeded
their ancient wisdom, i.e., their accumulated knowledge of the sea.
1
In cultures such as the Moken, and
0273-2297/$ - see front matter Ó2009 Elsevier Inc. All rights reserved.
doi:10.1016/j.dr.2009.04.001
*Corresponding author. Fax: +49 5121883479.
E-mail address: wgreve@uni-hildesheim.de (W. Greve).
1
For the general story see, e.g. http://www.cbsnews.com/stories/2005/03/18/60minutes/main681558.shtml (date of validation:
4th July 2008) or http://ngm.nationalgeographic.com/ngm/0504/feature4/online_extra.html (date of validation: 4th July 2008);
with particular respect to the wisdom of older Moken people, see, e.g., http://www.rense.com/general62/pretsn.htm (date of
validation: 4th July 2008).
Developmental Review 29 (2009) 163–179
Contents lists available at ScienceDirect
Developmental Review
journal homepage: www.elsevier.com/locate/dr
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surely for all of our ancestors during the greater part of human history, who cannot rely on written
texts, magnetic patterns on small discs, and other ways of conserving rare experiences, human mem-
ory is the only place to save such knowledge (Diamond, 2001). Thus, it may be useful, and sometimes
even necessary, to have people around who are old and sane enough to remember those valuable,
though seldom used, experiences and pieces of knowledge. In short, older folks are a ‘‘reservoir of fam-
ily wisdom” (Neugarten & Weinstein, 1964), and possibly ‘‘repositories of knowledge” (Lee, 1997) not
only for their relatives, but even for non-kin members of the social group they belong to.
Extending on this anecdotal evidence, we argue more generally that older people, and, therefore,
the human possibility to become old, afford real advantages, not only for aged people themselves,
but also for their children and grandchildren, their kin, and their neighbors. In the following, we pro-
pose that aging, irrespective of its obvious drawbacks, can be viewed as an evolutionary adaptation
(Mergler & Goldstein, 1983). More precisely, we argue that, while senescence may be an evolutionary
byproduct, longevity and in particular later periods of the lifespan are a result of positive natural selec-
tion (Carey & Judge, 2001). We attempt to demonstrate that several answers to the question of why we
do – or can – grow old (in particular, the so-called ‘‘grandmother” hypothesis) are, though partly sat-
isfying, not a sufficient explanation of longevity and, hence, need to be extended and enhanced by psy-
chological (developmental) perspectives. We argue, then, that old age, once evolved, became itself a
major force in human – in particular cultural – evolution, resulting in a mutual interdependence of
the evolution of longevity and culture.
The core of our argument focuses on those attributes that can, and usually do, accompany aging,
and particularly support the function of older people as providers of experience, knowledge, and wis-
dom. Starting from this point of view, we attempt to sketch an outline of an evolutionary psychology
theory of aging. In doing so, we argue that the emergent perspective of evolutionary developmental
psychology (e.g., Bjorklund, 1997; Bjorklund & Pellegrini, 2002; Burgess & MacDonald, 2005; Ellis &
Bjorklund, 2005; Geary & Bjorklund, 2000) can and should be extended to a lifespan perspective. In
other words, in addition to ‘‘the adaptive nature of immaturity” (Bjorklund & Green, 1992), we argue
in favor of ‘‘the adaptive value of periods beyond maturity”.
Towards an evolutionary developmental perspective: beyond immaturity
Evolutionary perspectives have taken hold within mainstream psychology within the past two dec-
ades (see Barkow, Cosmides, & Tooby, 1992; Buss, 2005; Daly & Wilson, 1988) and have been applied to a
wide range of topics including mating (e.g., Buss, 1999), helping (e.g., Nowak, 2006), homicide (e.g., Daly
& Wilson, 1988), language (Pinker, 1994), cognition (Barkow et al., 1992; Gigerenzer, 2000), education
(e.g., Geary, 2007), and family psychology (e.g., Salmon & Shackelford, 2007), among others (for compre-
hensive overviews, see Buss, 2005; Dunbar & Barrett, 2007). Among the most fascinating and unique fea-
tures of humans is the fact that we have a biography, that is, a structured development. Thus, the course
and structure of our ontogeny ask for an evolutionary explanation as do our brain or our social behavior.
This demand goes well beyond the ‘‘embryological” task of explaining the building of our body: we need
to explain the psychological structure of our life course. This explanation is the task for evolutionary
developmental psychology (Belsky, Steinberg, & Draper, 1991; Bjorklund, 1997;Bjorklund & Hernández
Blasi, 2005; Bjorklund & Pellegrini, 2002; Bogin, 1999; Burgess & MacDonald, 2005; Ellis & Bjorklund,
2005; Geary, 2006; Geary & Bjorklund, 2000).
The adaptive nature of immaturity: understanding early stages of development
Humans need a long time to reach adulthood, much more so than our primate cousins (Poirier &
Smith, 1974). Since it is rather dangerous to be young, risk prone, playful, and overconfident (i.e.,
immature; Bjorklund, 2007), several features and attributes of infancy and youth likely posses their
own adaptive value (Bjorklund & Pellegrini, 2002), since otherwise childhood and youth would have
been selected against. For instance, children’s play has been proposed to provide a host of benefits,
some immediate and some delayed, including regulating body temperature, establishing one’s posi-
164 W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179
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tion in the social hierarchy, tool use, and learning social skills necessary for intrasex competition and
cooperation, among others (Pellegrini & Bjorklund, 2004; Pellegrini & Smith, 1998).
Beyond maturity: the challenge of lifespan developmental psychology
Although these arguments provide a plausible explanation for our prolonged childhood and youth,
human development is more than maturation. According to the lifespan perspective on human devel-
opment, which was first articulated almost four decades ago (Baltes & Goulet, 1970; Baltes, Reese, &
Lipsitt, 1980), human development is characterized by plasticity and permanent change of the individ-
ual’s experiences of gains and losses throughout the entire lifespan (Baltes, 1987; for comprehensive
textbooks see, e.g., Bjorklund & Bee, 2008; Brandtstädter & Lindenberger, 2007; Feldman, 2006). There
is now abundant research demonstrating the fruitfulness of a lifespan perspective in developmental
psychology (e.g., Baltes, Lindenberger, & Staudinger, 2006; Baltes, Staudinger, & Lindenberger, 1999;
Staudinger & Lindenberger, 2003). However, given a 60-fold variation within mammals (2-year-mice
to 122-year humans; Carey, 2003; Kirkwood, 1999; Smith, 1993; for overview see Carey & Judge,
2000), our lifespan is a remarkable feature of our species. While our hominid ancestors reached a max-
imum lifespan of 40–50 years, comparable to that of our present-day primate cousins (Carey, 2003;
Judge & Carey, 2000), the extended life expectancy of subsequent members of the Homo clade has al-
most doubled since we took a separate evolutionary pathway. Only very few animals on earth attain a
longer lifespan (some mammals, e.g., elephants, and some non-mammals, e.g. turtles, sea cucumbers;
for overview Carey & Judge, 2000; Deevey, 1947; Gavrilov & Gavrilova, 1991), representing no homo-
geneous taxonomic group but rather specialized forms within heterogeneous groups (Smith, 1993).
Hence, we have to explain both its length and its structure from an evolutionary point of view.
Beyond longevity per se, particularly the emergence of periods of life beyond reproduction begs an
evolutionary explanation (Carey & Judge, 2001; Hawkes & Paine, 2006; Hrdy, 1999). In fact, the mere
existence of post-reproductive stages seems counterintuitive from an evolutionary perspective. If re-
sources are scarce (which is the presupposition of evolution to occur at all), the mere existence of non-
reproductive individuals (competing for food and safety) is a threat for the reproductive ones. More-
over, in social animals the aged often times require additional support from their grown-up children,
other relatives, or even unrelated group members who are still in the stage of reproduction, which, in
turn, may even endanger their primary ‘‘duty” to reproduce (Kaplan, 1997; Kaplan & Gangestad, 2005;
Kaplan, Hill, Lancaster, & Hurtado, 2000). In short: aging is costly, and, if taken in isolation, seems to be
maladaptive relative to its absence (Partridge, 1997). Why then do we age (Medawar, 1952)?
Several evolutionary theories of aging (e.g., Austad, 1997a; Finch, 1990; Rose, 1991) claim that
aging processes of decreasing gains and increasing losses and difficulties can be explained by a declin-
ing force of natural selection in post-reproductive stages (Baltes, 1997; Jazwinski, 1996; Kirkwood,
2003; Wachter, 1997; for possible genetic costs of aging see, e.g., Kirkwood & Holliday, 1979;
Westendorp & Kirkwood, 1998). If enough offspring are produced, so this argument goes, the longevity
of the parent is inconsequential from an evolutionary perspective. An illustrative example for this
argument is the expression of some genetic diseases that express beyond the age of reproduction
(Rose, 1991; Williams, 1957) such as Huntington’s disease (Austad, 1997a). The vast majority of bio-
logical theories of aging (for introductions see Austad, 1997a; Holliday, 2007,orKirkwood, 1999; for
comprehensive overviews see Charlesworth, 1980; Finch, 1990, 2007; Medvedev, 1990) deal with this
very question of why aging entails a decrease of competencies and resources. Following this line of
argument, Baltes (1997; Baltes et al., 1999, 2006) argued that the decreasing degree of adaptation
(i.e., the increasingly worse gain/loss ratio) has to be outbalanced by an increasing degree of cultural
compensation, which, in turn, may explain why aging is a historically young phenomenon. Yet, even if
this were true and the complete answer (which we doubt, as will become clear shortly), it still leaves
open the question of why and how longevity has occurred in the first place (Medvedev, 1990; Sacher,
1978). The question of why we don’t die after reproduction is far from trivial since many species have
followed this evolutionary pathway. The most often cited example of this ‘‘switch off”-model is the
salmon that dies just after releasing a large number of fertilized eggs (Finch, 1990). In fact, the mere
existence of menopause prows that a ‘‘switch off”-device is possible also for humans. Even somewhat
W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179 165
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more drastic, the females of some spider species kill and eat their mates once they have ‘‘done their
duty.”
2
A possible confusion: potential, not actual age is the issue
If we are to explain human longevity, it is important to emphasize that the possibility of growing old
is not an evolutionary ‘‘young” phenomenon, although the mean life expectancy has remarkably
increased during the last hundred years (Acsádi & Nemeskéri, 1970; Austad, 1997a; Gavrilov &
Gavrilova, 1991; Meier & Vaupel, 2003; Smith, 1993; Vaupel et al., 1998) as a result of enhanced
health services and medical techniques, improved nutrition, hygiene, and, in particular, reduced infant
mortality rate (Austad, 1997a). Yet, the maximum age has remained largely stable throughout the
history of our species
3
(Austad, 1997a; Cutler, 1975; Fries, 1980; Hawkes & Blurton Jones, 2005; Kaplan,
1997; Weiss, 1981; but see Carey, 2003; Wilmoth, 1997; Wilmoth & Lundstrom, 1996, with respect to
the last century). Although there is a certain degree of uncertainty here since many cases of extreme
longevity fail to be validly proven (Austad, 1997a; Christensen & Vaupel, 1996; Gavrilov & Gavrilova,
1991; Gavrilova & Gavrilov, 2005; Wilmoth & Lundstrom, 1996), and though the majority of humans
did not reach it until quite recently, a maximum age of about a 100 years has always been the possible
lifespan for humans.
4
Studies with contemporary hunter–gatherer societies demonstrate that even in
these societies women at the age of 45 can expect to live for another 20 years (e.g., Blurton Jones,
Hawkes, & O’Connell, 2002; Gage, McCullough, Weisz, Dutt, & Abelson, 1989; Hawkes & Paine, 2006;
Voland, Chasiotis, & Schiefenhövel, 2005a). Most probably, some individuals in all societies have attained
‘‘old age,” perhaps not reaching the species maximum, but far exceeding the average life expectancy for
their group. Actually, the percentage of individuals growing old beyond reproductivity has had to be rel-
atively large, since otherwise it would be impossible to find some adaptations in these later phases of life,
the most prominent of which is the phenomenon of menopause. Its evolution, of course, presupposes a
lifespan that is long enough to leave space for this interruption to occur. To be sure, the evolution of
menopause requires an evolutionary explanation itself (to which we will return shortly), but the mere
existence of it supports the argument that, in principle, ‘‘aging is old.” So, why can we age?
Towards an evolutionary theory of aging: grandmothers and beyond
One possible answer claims that aging is perhaps just a mere side effect (and, thus, a ‘‘spandrel”;
Gould & Lewontin, 1978) of being healthy during adulthood (like the ‘‘longevity” of a car beyond its
guaranteed age; Kirkwood & Rose, 1991; Smith, 1993; Vaupel, 1997). Yet, we (possibly) live beyond
our age of reproduction for several decades and against the tendency to accumulate physiological
and even genetic problems (Abrams, 1991; Kirkwood, 1977; Kirkwood & Holliday, 1979; Lithgow &
Kirkwood, 1996; Westendorp & Kirkwood, 1998) – which, unfortunately, most of our cars don’t. More-
over, our longevity entails maintenance procedures both on the individual (e.g., DNA repair) and social
levels (Baltes, 1997; Kaplan et al., 2000; Voland et al., 2005a). Hence, the phenomenon of staying alive
beyond reproduction has to be outbalanced by its evolutionary advantages in terms of fitness, other-
wise longevity would not have been retained but rather eliminated by natural selection.
2
Some authors have claimed that sometimes humans show this kind of behavior (i.e., killing the elderly; Simmons, 1945). The
evidence, however, is scarce, and the circumstances, if such behaviors are to be observed, have to be extraordinary.
3
Though longevity probably has changed within the homo clade from Australopiethcus to Homo sapiens (Carey & Judge, 2001).
4
Actually, a little beyond that (Robine & Allard, 1998). There is a recent increase of so-called ‘‘supercentenarians” (persons
reaching the age of 110 years or more) since the mid-1970s (Robine & Vaupel, 2001), and the best confirmed report of a
supercentenarian is Madame Calment who died at an age of 122 (e.g., Austad, 1997a). Perhaps, our lifespan maximum is a little bit
higher than previously expected, but wherever it is, it is close to that number (which is different from the – much lower: 85 – life
expectancy at birth; Olshansky, Carnes, & Cassel, 1990). We do not want to delve into the debate of whether there is a ‘‘sharp” limit
for longevity (Carey, 2003); most probably, aging and dying is not simply genetically programmed (Austad, 1997a; Holliday, 2007;
for a detailed discussion see, e.g., Kanungo, 1994). Yet, wherever the bounds are, we stick to the conviction that we are destined to
die, sooner or (somewhat) later.
166 W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179
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The riddle of menopause: the grandmother effect
Human females experience their menopause well before other physiological signs of senescence
become apparent, and live, roughly, one-third of their lives as post-reproductives, which is unusual
among mammals in general and primates in particular (Austad, 1997a, 1997b; Caro et al., 1995;
Hawkes & Blurton Jones, 2005; Sorensen Jamison, Jamison, & Cornell, 2005).
5
Obviously, the occurrence
of menopause must have special advantages (Lancaster & King, 1985; Pavelka & Ferdigan, 1991; Peccei,
1995, 2000, 2001, 2005; Rogers, 1993; Sherman, 1998; Turke, 1997; Williams, 1957; for an introduction,
see Voland, Chasiotis, & Schiefenhövel, 2005b). The main line of argument here is referred to as the
grandmother effect (Hawkes, 2003, 2004; Hawkes & Blurton Jones, 2005; Hawkes, O’Connell, Blurton
Jones, Alvarez, & Charnov, 1998; Hawkes, O’Connell, & Blurton-Jones, 1997; Hill, 1991; O’Connell,
Hawkes, & Blurton-Jones, 1999; Sorensen Jamison, Jamison, Cornell, & Nakazato, 2002; Sorensen Jamison
et al., 2005; for earlier attempts see Hrdy, 1981; Williams, 1957; for overview, see Euler & Michalski,
2008; Voland et al., 2005b), which proposes that post-menopausal females contribute to the success
and survival of the children of their children (Alvarez, 2000). This argument rests on the insight that
any evolutionary success depends on the ability of one’s offspring to reproduce themselves. Beyond
being a relief for the mother, a healthy and wise grandmother may provide necessary support in raising
offspring, for example, during the period of pregnancy when the earlier children are still young (Smith,
1991). Remember that humans have an extended phase of immaturity and thus need of adult assistance,
care, and protection even after they have been weaned. Consequently, Hrdy (1999) argued that the evo-
lution of longevity and of an extended period of immaturity are related; the advantages of immaturity
fuel a prolongation of these periods, but, at the same time, require longer periods of care from adults.
There is growing evidence that granny’s ‘‘help at the nest” enhances the reproductive success of her
offspring by enabling them to attain a higher birth rate (Mace & Sear, 2005) and increasing the prob-
ability of survival of their grandchildren (e.g., Sorensen Jamison et al., 2005). Evidence stems both from
historical data (e.g., Beise, 2005; Beise & Voland, 2002; Lahdenperä, Lummaa, Helle, Tremblay, &
Russell, 2004; Sorensen Jamison et al., 2002; Voland & Beise, 2002) and from anthropological studies
investigating contemporary hunter–gatherer societies such as the Hazda of northern Tanzania and the
Ache of eastern Paraguay, among others (e.g., Hawkes, O’Connell, Blurton Jones, Alvarez, & Charnov,
1998; Kaplan et al., 2000; Mace & Sear, 2005; Sear, Mace, & McGregor, 2000, 2003; Sorensen Jamison
et al., 2002). Throughout our species’ history, grandmothers have fulfilled a great variety of roles and
functions (Bengtson & Robertson, 1985; Brake & Büchner, 2007; Johnson, 1983; Kahana & Kahana,
1971; Robertson, 1977; Schölmerich, Leyendecker, Citlak, Miller, & Harwood, 2005; Smith, 1991). They
not only provide food for their adult children’s families, but, in particular, provide experience and
knowledge during risky periods in their grandchildren’s lives (e.g., weaning and transition to milk sub-
stitute; Beise, 2005). Grandmother’s role as a competent (experienced) birth attendant might have
been of additional importance. Several studies show that the helping grandmother can be identified
even under modern social conditions (Brake & Büchner, 2007; Denham & Smith, 1989; Kivnick,
1983; Nosaka & Chasiotis, 2005; Schölmerich et al., 2005; Smith, 1991).
However, things may be more complicated than they seem at first glance (Euler & Michalski, 2008;
Voland et al., 2005a). For instance, since fatherhood is uncertain whereas motherhood is not, it could
be expected that it is the maternal grandmothers in particular who contribute to the welfare of their
children and grandchildren. For example, in contemporary rural Ethiopia, the assistance of a maternal
grandmother is associated with higher child survival rates (Gibson & Mace, 2005), and the presence of
a maternal grandmother is related to higher fertility and survival rates for Canadian and Finnish farm
families (Lahdenperä et al., 2004). A similar story is seen from historical records. Based on more than
150 years of German births (1720–1874), children without a living maternal grandmother were more
likely to die than those without a living paternal grandmother. In some age intervals, the difference in
death rate was as large as 60% (Beise & Voland, 2002). On the other hand, paternal uncertainty could
5
However, there is some controversy among biologists here. Among wild mammals, only pilot whales and elephants seem to
experience menopause (Carey & Gruenfelder, 1997; Whitehead & Mann, 2000), although other research indicates that rhesus
monkeys and gorillas in captivity also experience an extended post-reproductive period (e.g., Atsalis & Margulis, 2006; Cohen
(2004), Walker, 1995) argues that menopause is a more general trait.
W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179 167
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also explain why the social role of the paternal mother-in-law has always been described as suppress-
ing for the daughter-in-law (Voland & Beise, 2005). Actually, there is some evidence that this problem
influences the advantageous role of grandmothers (for an overview see, e.g., Euler & Michalski, 2008;
Mace & Sear, 2005; but see Leonetti, Nath, Hemam, & Neill, 2005). For instance, recent data from a
study with the Dogon of Mali suggest that a longer survival of paternal grandmothers may even be
detrimental both for growth rates and survival rates of grandchildren (Strassmann & Welch, 2007).
However, as Beise (2005) aptly remarks, many woman have both adult daughters and sons, so they
are maternal and paternal grandmothers at the same time. Moreover, the grandmother effect, even
if qualified by lineage, seems to depend on further circumstances, such as patrilocal vs. matrilocal
societies, to name just one factor (Mace & Sear, 2005; Voland et al., 2005b).
Beyond ‘‘help at the nest: the adaptive value of cumulative knowledge and experiences
In fact, the grandmother effect cannot be the sole explanation of longevity because men also grow
old. Though grandfathers do not experience a physiological menopause, almost all of them show – or
experience – a behaviorally post-reproductive period of their life as well (Austad, 1997b; Carey &
Judge, 2001). Thus, the problem of why males also age doesn’t differ fundamentally from the problem
of post-menopausal females. Actually, there is some evidence that aging males (and their mating pat-
terns) are part of an evolutionary explanation of human lifespan extension (Tuljapurkar, Puleston, &
Gurven, 2007). Thus, it is still unclear whether the evolution of longevity has been a male-driven or
a female-driven process (Voland et al., 2005a): Are the fitness gains of longevity produced by long-
er-active male hunters (Kaplan et al., 2000) or by female gatherers (Hawkes & Blurton Jones,
2005)? However, old men in traditional societies seldom, if ever, take care of babies. Though there
are only a few studies focusing on the role and function of grandfathers (e.g., Kivett, 1985, 1991; Radin,
Oyserman, & Benn, 1991; for a recent overview see Brake & Büchner, 2007), recent data suggest that a
grandfather effect in the evolution of human lifespan seems to be negligible (Lahdenperä, Russell, &
Lummaa, 2007).
Throughout human history, social institutions such as the Roman ‘‘Senate” (meaning: ‘assembly of
old men’) reflect the insight that cumulated experience of its older members is a valuable asset for a
group or society. In all human societies, in particular preliterate societies (a state characterizing the
vast majority of human history), the preservation of knowledge depended almost completely on the
memories of experienced (and sufficiently sane) old people (Diamond, 2001; Mergler & Goldstein,
1983). To have them around, to have access to their experiences, knowledge, and thus their advise,
helped the family, kin, and the social group to which they belonged in their struggle for life and repro-
duction (Carey & Gruenfelder, 1997). We propose to name this particular fitness-enhancing effect of
the cumulative and integrated knowledge (‘‘wisdom”) of old members of a group as the Nestor effect.
In Homer’s Iliad, Nestor, the oldest member of the group of Greek kings at the siege of Troy, was re-
spected and well liked by the other Greek leaders, though (or because) he tended to make rather
long-winded speeches and liked to tell long stories about his past exploits. His name has become pro-
verbial for an old and respected advisor.
The risky behavior of children requires mature persons who take care of them. In other words, the
adaptive value of immaturity depends, to a certain degree, on the presence of mature and experienced
caretakers. Particularly in times of need, such as droughts, famines, or cold spells, humans often do
need sage advise from experienced (and hence older) individuals who may remember that it has
proved useful to search for mammoths in some distant valley, that water still can be found in a
particular spring, that using buffalo fat as an additional layer between skin and clothes may prevent
severe frostbites, or that specific changes in the sea foretell of the occurrence of a dangerous tsunami.
This argument presupposes, of course, that humans (brains in general, for that matter; Finch &
Ricklefs, 1991) are capable of retaining such old memories. Several studies indicate that, whereas
the mechanics of cognitive functions (such as speed of information processing and working-memory
span) decrease with age, the pragmatics, in particular cumulative knowledge and experience, do not
show a negative trend in age (Baltes et al., 1999, 2006). In particular, the fact that those contents of our
long-term memory that refer to our youth are less prone to be forgotten if we are old (Rubin, Wetzler,
168 W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179
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& Nebes, 1986) fits well with the claim that exactly those memories are the most valuable ‘‘capital” of
older people (Carey & Judge, 2001; Mergler & Goldstein, 1983). All members of a group experienced
and likely have memories for recent events. Thus, the ‘‘oldest” memories are most probably those that
make the memory of the Nestor unique, as it were.
The value of older people as Nestors entails the maintenance of the ability to share their valuable
memories with the younger members of their kin or group. Tyszkova (1991) demonstrated that con-
versation is the most frequently used category of activities between grandparents and grandchildren.
In fact, older people often remain fully capable of social interactions even after cognitive deficiencies
arise (Carstensen & Charles, 1998, 2003). It may be added here that the elderly are by no means nec-
essarily the weakest, frailest, and unhealthiest of their birth cohort. To the contrary, very old persons
are pretty untypical persons with respect both to their vulnerability and their risk-proneness (e.g.,
Crimmins, 2001), which most probably contributes to their suitability as Nestors.
There is evidence from both traditional and modern cultures that the presence of older people in a
group provides valuable assistance to younger members, for instance with respect to food and labor
(e.g., Blurton Jones, Hawkes, & O’Connell, 1996, 1999; Hawkes, O’Connell, & Blurton-Jones, 1997; Kap-
lan et al., 2000; Sear et al., 2003) or caring and solicitude (e.g., Euler & Weitzel, 1996; Gaulin, McBur-
ney, & Brakeman-Wartell, 1997; for cross-cultural comparisons see Amoss & Harrell, 1981).
Furthermore, there is also some evidence that something akin to the Nestor effect is found in some
non-human species: elder individuals play important roles, for instance, in group cohesion, as caregiv-
ers, guardians, leaders, or teachers (Carey & Gruenfelder, 1997). One prominent example is elephants
(Buss, 1990; Carey & Gruenfelder, 1997; Eltingham, 1982; McComb, Moss, Durant, Baker, & Sayialel,
2001). Like apes and humans, they have an extended juvenile period, live in socially complex groups,
and are long-lived, with females living a decade or so beyond producing their last viable egg (Hart,
Hart, & Printer-Wollman, 2008). Elephants’ social groups are lead by the oldest female (Buss, 1990;
Eltingham, 1982). McComb et al. (2001) reported that age of the matriarch was positively associated
with her family’s reproductive success, probably ‘‘through its effects on the acquisition of social
knowledge” (p. 493f.). There is some anecdotal evidence that clans led by older matriarchs do better
in coping with severe droughts, e.g., since the matriarch remembers the location of a certain water
source (Foley, 2002; Payne, 1998; cf. also Carey & Gruenfelder, 1997). Likewise, hamadryas baboons
benefit from expertise of some elder members in their troops (Kummer, 1995, see in particular p.
237ff.; for a more general approach see Avital & Jablonka, 2000). Even among whales, older individuals
may take over valuable social functions within a social group (Carey & Gruenfelder, 1997).
Following this line of argument, it should be possible to identify selective conditions that may favor
the evolution of longevity, such as unpredictable or scarce environments, extended parental care for
few offspring, and, in particular, social cooperation and communication (Greve, Kierdorf, & Kierdorf,
2009). Hence, one empirical line starting from our claim should ask for the evolutionary advantages
of older group members, particularly in these environments (comparison of groups with and without
Nestors).
Beyond generativity: making (individual) sense makes (evolutionary) sense
There is no doubt, however, that the human lifespan, both with respect to its duration and struc-
ture, is as unique as our species is in general. Thus, our argument goes beyond the adaptive value of
cumulative experiences. Humans’ possess unique competencies to process our stored knowledge: We
connect it, integrate it, evaluate it, and thus form a coherent whole of it. The most mature form of this
kind of integration has been named, from ancient times until today, as ‘‘wisdom”. Wisdom includes,
beyond the integration of knowledge and experiences of a whole lifespan (including knowledge ac-
quired from other members of the society), procedural knowledge about where to get information
and how best to give advice, lifespan contextualism about the person’s age and station in life, and
understanding that back-up plans may be necessary (Smith & Baltes, 1990; Sternberg, 1990). In short,
wisdom entails ‘‘a well-balanced coordination of emotions, motivation, and thought, with good judg-
ment and the ability to offer advice in difficult and uncertain matters of life” (Staudinger & Dörner,
2006, p. 674). The point of argument here is that the ability to integrate accumulated experiences into
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a cognitive network that makes this information both valuable and available has significant fitness
advantages beyond mere storage. This holds in particular given the fact that wisdom is, to a large
extent, interactive by nature (Baltes & Staudinger, 1996). However, the emergence of wisdom is asso-
ciated with age (Baltes & Staudinger, 2000; Staudinger, 2000, 2001, 2005; Staudinger & Dörner, 2006;
Staudinger, Dörner, & Mickler, 2005).
There is an interesting self-enhancing dynamic here, in that self-related wisdom (Staudinger, 2001;
Staudinger et al., 2005) includes processes that help to keep the aging mind stable, sane, and, thus,
valuable for others. A large body of research shows that, although the ratio of gains to losses becomes
increasingly unfavorable with age both objectively (Baltes et al., 2006) and subjectively (Heckhausen,
Dixon, & Baltes, 1989) for the large majority of people, subjective well being does not decrease in later
adulthood (Brandtstädter, Wentura, & Greve, 1993; Staudinger, 2000). This evinces the existence of
various mechanisms at work, which buffer or completely absorb the impact of increasingly negative
developmental influences (Greve & Staudinger, 2006).
This approach fits perfectly with Erikson’s (1950, 1959/1980; 1997; Erikson, Erikson, & Kivnick,
1986) theory of lifespan development, in that his approach also encompasses a life phase beyond gen-
erativity – integrity. The achievement of integrity not only prevents individual declines in well being
(‘‘despair”), but, from an evolutionary point of view, this capacity is exactly what makes age useful and
adaptive. To have at least some Nestors available in a family or group who possess information that
the family may need from time to time, would obviously provide a measurable fitness advantage
for its members.
Beyond good advice: when grandparents become grantparents
Perhaps it is more than just an etymological footnote that the very concept of ‘‘inheritance” is used
primarily to refer to the transmission of ontogenetically acquired (accumulated) advantages (i.e.,
material, social, or cognitive goods, e.g., money, status, ‘‘rules to live by,” or recipes). Thus, it is ‘‘genetic
inheritance,” rather than ‘‘cultural inheritance,” that is the metaphorical use of the inheritance con-
cept. Actually, beside our genes we inherit from our ancestors cultural goods such as a house, money,
or works of art, but also more abstract rights and obligations, such as debts, credits, and rights (for
instance, the right to be king or queen), and, above all, knowledge (Bjorklund, 2006). In fact, it is an
important function of grandparents, at least in many cases, to transmit valuable goods and cultural
items (including non-material ones) to the next generations (Carey & Judge, 2001; Goodnow & Law-
rence, 2008). The transfer flows from the elderly out weigh, as a rule, the transfer flows to them (Lee,
1997) both in traditional (e.g., Kaplan, 1994) and modern societies (e.g., Brake & Büchner, 2007; Judge
& Hrdy, 1992; Kaplan, 1996;Lee, 1997, 2003). Moreover, material inheritance increases the probabil-
ity of having grandchildren (Lynch, 1996). Though this effect has increased in modern societies (be-
cause the benefits from inheritance are privatized, whereas the financial burdens of caring for elder
members of the society [pensions, health care] are socially distributed), it can be assumed that the
evolutionary advantages of ‘‘real inheritance” were even more important in the dawning of human-
kind. A finely honed stone axe, for example, was likely a valuable grant, since its manufacture was dif-
ficult and energy consuming, and its possession afforded the user access to other important resources
(e.g., food, status, an impressed future mate). Moreover, its personal receipt from an ancestor, includ-
ing some advise about how to use it properly, may extend even beyond its practical use and also in-
clude additional cultural or symbolic value (e.g., ‘‘the” axe of the village: s/he who owns it is destined
to be the leader – as in the case of the holy sword of England, ‘‘Excalibur”).
Aging and culture: dynamics of reciprocal enhancement
Baltes (1997, Baltes et al., 1999) argued that age is in need of cultural compensation for its inherent
losses and weakness. We do not doubt that, in many cases, older people do need support from younger
persons and thus cultural compensation. In fact, our argument, to a certain degree, started from the
acceptance of this necessity. However, beyond pointing to the fact that the mere possibility of growing
old still begs for an evolutionary explanation, we would like to add the complementary perspective,
170 W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179
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according to which the existence of old age is, at the same time, an important factor for the historical
emergence of culture (Carey & Judge, 2001). Culture depends on memory in the first place: valuable
information can only survive if there is a place in which it can ‘‘exist.” This holds in particular for expe-
riential (episodic) knowledge, such as knowledge about useful plants (against rare diseases), useful
knowledge (indicators of rare crises such as floods), useful strategies (against rare problems such as
unusual droughts), or useful techniques (e.g., of hunting rare prey). Moreover, old age entails not only
having more material to think about, so to speak, but also having more time to do so. As obligations of
mature adulthood subside, the elderly have increasing ‘‘leisure” time to devote to other activities.
Thus, the emergence of longevity may be the starting point, possibly even the necessary precondition,
for the evolution of advanced culture, that is, for the evolution of a new evolutionary dimension (Jab-
lonka & Lamb, 1995, 2005).
6
Culture, so to speak, is a pool of knowledge, stored in the brains of a pop-
ulation of people (Boyd & Richerson, 2005, p. 6). Because cultural evolution rests to a large degree on
normative and instructive communication (Boyd & Richerson, 2005; Tomasello, 1999), the existence,
and capability of Nestors may be one essential condition for culture to evolve.
The productive loop: Nestors produce culture that, in turn, secures their existence
Of course, the emergence of culture is a rather complex phenomenon (Boyd & Richerson, 1985,
2005; Durham, 1991; Tomasello, 1999), and we do not wish to claim that aging is the decisive factor
for its explanation. However, it may be worth thinking of it as an important factor fueling, accelerat-
ing, and, in particular, stabilizing the emergence of culture.
Our central argument is an interactive, bidirectional one. Certainly, we do not doubt that culture
also supports aging (Baltes, 1997), but we want to stress that, at the same time, aging (accumulating
pragmatics of life) supports, or even affords, culture. The presence of some old and wise persons
(‘‘back-up copies,” ‘‘integrators,” etc.) enhances fitness, at least of those older person’s kin, but prob-
ably far beyond the bounds of the family. At the same time, because these elders have valuable knowl-
edge, the intellectual capacities, and ‘‘free” time, they are able to invent, refine, or propagate basic
tools of culture, perhaps painting, stone axes, letters, or political protocols. Although nursing young
mothers and hunting and fighting young men surely contributed to the development of ‘‘culture,” they
likely neither had the time nor experience to make great headway in cultural maintenance alone.
Rather, it seems reasonable to assume that older members of the group likely played a prominent role
in the preservation and promotion of culture (Carey & Judge, 2001). Beyond technical or scientific
inventions and innovations, culture essentially consists in progress of humanity, that is, in perceiving
and regarding other humans’ needs. Moreover, culture entails traditions of shared knowledge far be-
yond technical skills (e.g., fairy tales, songs, customs, etc.) that create and enhance the sense of com-
munity. In this respect, it seems plausible to assume that the elder members of a group or society soon
became important caretakers of culture.
This argument can be taken one step farther. Once cultural traditions and social norms are estab-
lished, it is probable, perhaps necessary, to install some self-serving ‘‘memes” (Dawkins, 1976), that is,
rules that promote the care and respect for the elderly, who are the receptacles of these capabilities.
The more of these rules and norms that are in force, the more older persons will survive, perhaps more
than are necessary for the original ‘‘purpose,” so there will be more resources available to establish,
maintain, and modify culture (liberal arts, useful inventions, etc.), which in turn will foster social
norms encouraging a culture of respect for the aged (for discussion of these self-sustaining ‘‘memet-
ical” dynamics, see Aunger, 2002; Blackmore, 1999; Distin, 2004). One likely driving force of this dy-
namic interaction was the invention and preservation of medicinal knowledge and skills, which serves
both younger and older members of the family, tribe, or society. In short, aging supports culture, and
culture supports aging. If this argument proves to be tenable, it may shed new light on the evolution-
6
We do not want to delve into the discussion here of whether there is culture in animals other than humans (e.g., chimpanzees:
Whiten, 2005; Whiten et al., 1999; orangutans: van Schaik et al., 2003; whales: Rendell & Whitehead, 2001; even guppies,
Dugatkin, 2001). There may be aspects of informational transmission by social learning in nonhuman species, but by any means it
is clear that only human culture has created a special dynamic on its own – a new dimension of development (Jablonka & Lamb,
2005; Tomasello, 1999).
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ary biology of aging. Perhaps the traditional argument that age-related declines are inevitable (argu-
ably as the consequence of decreasing selective pressure; Rose, 1991) isn’t evolutionary enough, as it
were.
The evolutionary interdependence of culture and aging among humans
Evolution never comes to an end. Once the Nestors, and hence experience, even wisdom, had
evolved, this established a new facet of the environment, resulting in a new configuration of selective
pressure. The wisdom of the Nestors created foreknowledge, prevention, and possibly cures with re-
spect to the circumstances that threatened their very existence, including inventions of how to ease
the burden to nourish the Nestors, eventually resulting in today’s medical research on Alzheimer’s
and other age-related problems and diseases.
Of course, this problem is much more complicated. Single factors, even those that seem substan-
tially correlated with aging (e.g., ‘‘encephalization”: the proportion of brain weight to body weight;
Hammer & Foley, 1996) cannot be viewed as explanations (there are, for instance, substantial differ-
ences in longevity between some new-world monkeys irrespective of the similarity of their respective
index of encephalization; Smith, 1993). The very same problem occurs for several other variables that
covary with maximum life span (e.g., metabolic rate, hibernation; de Magalhaes, 2008; Gavrilov &
Gavrilova, 1991) and that show several exceptions (bats, for instance, are a small species but outlive
many larger species). Moreover, there are numerous environmental factors that contribute to the sur-
vival rate of individuals in a population such as droughts, floods, storms, and predators, among them,
in particular, viruses and germs (Diamond, 1999; Ewald, 1993). Moreover, because these factors inter-
act with the Nestor effect (because one particular advantage of senior advisors is their very compe-
tence to deal with these threats and problems), there is no simple calculation of the result.
However, though this makes things difficult, it does not hurt the basic line of our argument. Given
a certain environmental constellation, a sufficient number of wise old persons is an adaptation. These
senior advisors, in turn, will change this very constellation, thus creating a new state of affairs, in that
an increase of the number of Nestors might have resulted in the creation of (more) culture, which en-
tailed capabilities to further increase the number of Nestors. That is, cultural changes brought about
by the presence of a sufficient number of Nestors not only served to compensate for the lack of bio-
logical support, but also to ease the burden older people represented for the group. In short: human
longevity may be a self-reinforcing process (Carey & Judge, 2001). Though a chicken–egg debate surely
is of no use here, we tend to concur with Carey and Gruenfelder’s (1997) claim that extended longevity
may even be viewed as a preadaptation for eusociality. The high level of maximum lifespan potential,
as compared to other mammals, surely represents a significant characteristic of humans and other pri-
mates (Cutler, 1976), and it is highly plausible to assume that it has contributed to their evolutionary
success (so far). For instance, the increase of brain size and the increase of lifespan may be linked in
vertebrate evolution (Holliday, 2007; Sacher, 1978; but see Gaillard, Allaine, Pontier, Yoccoz, &
Promislow, 1994).
Outlines of an evolutionary lifespan developmental psychology: perspectives for theory and
research
If the arguments outlined in this paper have not gone entirely wrong, the phylogenetic occurrence
of age (and aging) in Homo sapiens, as well as its shape and function, is understandable from an evo-
lutionary point of view. Though the core of our evolutionary argument is psychological by nature, we
do not intend to argue in favor of an alternative to biological approaches to the explanation of aging
(Austad, 1997a; Rose, 1991) or to deny their relevance. Instead, we argue that the psychology of aging
may add interesting perspectives to this emergent picture. It is a combination of psychological com-
petencies and abilities that, growing with age, make older people valuable for their kin. Older mem-
bers of a family or group provide useful or even necessary support for the younger members,
sometimes highly specialized (as in the grandmother case), sometimes more general (as in the tsu-
nami case). The cumulative knowledge and experience ‘‘stored” in the memories of older persons thus
172 W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179
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increase the fitness of the family that cares for their Nestors. Hence, there is positive selection for the
possibility of getting old: the principal reachable age is expected to extend, provided that the mind
and memory of (potential) Nestors is sufficiently capable of saving cultural expertise and knowledge
and sharing it with the descendants.
However, the very existence of older persons with wisdom and time on their hands alters the social
environment and thus the selective landscape and, finally, the course of evolution. In particular, the
Nestors may have played a vital role in the creation of a culture that supports caring for the elderly.
This in turn enhances the opportunities and resources not only for the family in general but also for
aged persons in particular, and thus the probability of getting old. As a consequence, culture and
old age mutually enhance themselves in a process of reciprocal reinforcement. In sum: the Nestor
effect may provide another example of what Baltes, Rösler, and Reuter-Lorenz (2006) called ‘‘bio-cul-
tural-co-constructivism,” perhaps a prototypical one. We argue that those families that (by random
variation) happened to produce members who lived longer, immediately experienced significant fit-
ness advantages (i.e., a greater number of surviving grand-offspring). In short, we claim that longevity,
and hence aging, is an adaptation. More precisely, we argue that only a certain way of aging (increas-
ing ability to integrate knowledge and experiences, increasing pragmatics of life, maintaining of long-
term memory contents until very old age, maintaining social competencies and motives, etc.) provides
these adaptive values of longevity.
Note at this point that our argument does not rest on the existence nor on the effectiveness of
group selection. We do not want to conceal that we sympathize with this concept (for an extended
discussion see Sober & Wilson, 1998; Wilson & Sober, 1994; Wilson & Wilson, 2007), and our argu-
ment would surely be further supported if group selection were proven to be a sound concept (pos-
sibly, our argument, given its credibility, may also conversely add support to the idea of group
selection). However, the core of our argument can rest on kin selection (i.e., inclusive fitness; Hamil-
ton, 1964) alone.
Two central conclusions can be drawn, should the ideas proposed in this paper prove to be sound.
First, our arguments brought forward in support of the Nestor Effect underscore the fruitfulness (and,
hence, the legitimization, as it were) of a lifespan perspective on human development. At first glance, a
evolutionary point of view may question this extended understanding of ‘‘development,” arguing that
the maturation of the zygote to an adult individual that is able to reproduce has to be conceptually
separated from the learning processes of this very adult. However, if later periods (life stages) do have
important evolutionary (i.e., reproductive) functions as well, a lifespan view on human development
seems biologically plausible. If the whole life history is an evolutionary product, and if also later stages
are reproductively important, a lifespan view of development is useful, or even necessary. Hence, our
arguments not only extend an evolutionary developmental approach to a lifespan perspective, but, at
the same time, support this approach against a more restricted concept of development
(‘‘maturation”).
Second, our arguments may fuel further ideas. In fact, several empirically testable hypotheses are
entailed in what we have sketched. In particular, fitness differences between (comparable) popula-
tions or groups are expected to covary with the number of Nestors as well with their ‘‘quality” (e.g.,
rate of dementia). There is some evidence indicating that the benefit of having Nestors around may
be even somewhat more tangible. Lee (1997, p. 217ff.) discusses a study that demonstrates that the
losses of farm profits due to bad weather conditions (statistically) depend on the present of older per-
sons. Likewise, the number of Nestors in a certain group may predict the enlargement of the mean lon-
gevity of that group in the next generation(s).
Moreover, the perspective we have argued for could provide theoretical feedback to evolutionary
developmental biology, in particular to life-cycle approaches (Bogin, 1999; Bogin & Smith, 1996; Bon-
ner, 1993; Hawkes & Paine, 2006; Raff, 1996). These approaches usually, if not always, focus on the
embryological point of view, that is, mainly looking at the earlier states in life. Our arguments suggest
that it may be worthwhile to look also at the later years of life, particularly at the post-reproductive
period(s). Here, several possible hypotheses can be investigated with respect to non-human mammals
as well (Greve et al., 2009).
The value of grandparents, in particular their culture-fueling significance, means, however, that
intergenerational communicative interaction works well. If generativity also entails the transmission
W. Greve, D.F. Bjorklund / Developmental Review 29 (2009) 163–179 173
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of knowledge, skills, and experiences across generations, we need to know much more about path-
ways and constraints of communication between generations. This area of research has been explored
only recently (e.g., Brake & Büchner, 2007; Kessler & Staudinger, 2007; Schölmerich et al., 2005; Smith,
1991), and we still know very little about the means of cultural generativity.
Finally, the lifespan approach to developmental psychology comprises more than claims about the
gains in addition to, and partly as compensation of, the losses of old age. In particular, one of the cen-
tral tenets of this approach is the multidirectionality and the plasticity throughout the lifespan (Baltes,
1987, 1997; Baltes et al., 1999). Though the arguments presented herein may contribute to an evolu-
tionary theory of human aging, they still largely refer to the ‘‘reliable reproduction” metaphor. How-
ever, a more open-structured theory of development that is compatible with evolutionary arguments
and insights, while at the same time compatible with tenets of plasticity and multidirectionality, is
obviously called for. How can an action-theoretical approach to human development and the idea
of intentional self-development (Brandtstädter, 2006; Brandtstädter & Lerner, 1999; Greve, Rother-
mund, & Wentura, 2005) be reconciled with the demands of reliability of reproduction of advanta-
geous attributes across generation (Greve & Ebner, 2007)? Further thinking is needed.
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