Mark,

I started to read this, and make some comments. The main question that occurred to me 
is why don't the authors consider such things as digging a burrow or migrating to 
avoid cold as genetically based instincts, as essentially phenotypical expression of 
genotype.  Isn't there such a thing as instinctive animal BEHAVIOR ?  If so isn't a 
form of phenotype  ( the paper mentions Dawkins' "extended phenotype")? Finally, why 
do they treat it as if it is incipient animal culture ?  Are they saying that child 
animals learn , for example, seasonal migration by imitating parents , and that it is 
not genetically based instinct ?

I make a couple of interjections in the excerpted text below, at points in their 
discussion where these questions occurred to me.

>>> [EMAIL PROTECTED] 08/21/00 07:34AM >>>

Niche Construction, Biological Evolution and Cultural Change.

1.0 AN EVOLUTIONARY FRAMEWORK FOR THE HUMAN SCIENCES

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1.1 Niche Construction

Building on ideas initially developed by Lewontin (1983), we have previously
proposed that biological evolution depends not only on natural selection and
genetic inheritance, but also on "niche construction" (Odling-Smee, 1988,
Odling-Smee, et al., 1996; Laland et al., 1996a). By niche construction we
refer to the same processes that Jones et al. (1997) call "ecosystem
engineering". Niche construction refers to the activities, choices and
metabolic processes of organisms, through which they define, choose, modify
and partly create their own niches1. For instance, to varying degrees,
organisms choose their own habitats, mates, and resources and construct
important components of their local environments such as nests, holes,
burrows, paths, webs, dams, and chemical environments. Many organisms also
partly destroy their habitats, through stripping them of valuable resources,
or building up detritus, processes we refer to as negative niche
construction. In addition, organisms may niche construct in ways that
counteract natural selection, for example by digging a burrow or migrating
to avoid the cold, 

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CB: Isn't this niche construction instinctive and a phenotype ?

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There are numerous examples of organisms choosing or changing their
habitats, or of constructing artefacts, leading to an evolutionary response
(Odling-Smee et al., 1996; Laland et al., 1996a). For instance, spiders
construct webs which have led to the subsequent evolution of various
camouflage, protection and communication behaviours on the web (Edmunds,
1974; Preston-Mafham & Preston-Mafham, 1996).

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CB: Are they saying this web construction or the subsequent camouflage, protection and 
communication behaviours are instintive or incipiently "cultural"  ? Are they passed 
on through genes or as a "tradition" ? I would assume the former.

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Similarly, ants, bees, wasps
and termites, construct nests which often themselves become the source of
selection for many nest regulatory, maintenance and defence behaviour
patterns. Many ant and termite species regulate temperature by plugging nest
entrances at night, or in the cold, by adjusting the height or shape of
their mounds to optimise the intake of the sun's rays, or by carrying their
brood around their nest to the place with the optimal temperature and
humidity for the brood's development (Frisch, 1975; Hansell, 1984). The
construction of artefacts is equally common among vertebrates. Many mammals
(including badgers, gophers, ground squirrels, hedgehogs, marmots,
monotrema, moles, mole rats, opossum, prairie dogs, rabbits and rats)
construct burrow systems, some with underground passages, interconnected
chambers, and multiple entrances (Nowak, 1991). Here too there is evidence
that burrow defence, maintenance, and regulation behaviours have evolved in
response to selection pressures that were initiated by the construction of
the burrow (Nowak, 1991). In all of the above examples there is often strong
comparative evidence suggesting that nest building is ancestral to the nest
elaboration, defence and regulatory behaviour (Hassell, 1984; Nowak, 1991;
Preston-Mafham & Preston-Mafham, 1996).

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CB: They have evolved, but how are they passed on to the next generation ? Genetically 
or as a "tradition" or "custom" ?

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Most cases of niche construction, however, do not involve the building of
artefacts, but merely the selection or modification of habitats
(Odling-Smee, 1988). For instance, many insects choose particular host

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CB: They "choose" it ?  Is this instinctive choice or "conscious" choice ?

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Niche constructing organisms may also substantially modify the environment
of their offspring, and even more distant descendants. Thus generations of
organisms not only inherit genes from their ancestors, but also a legacy of
natural selection pressures which have been modified by ancestral niche
construction. This legacy of modified selection pressures has previously
been labelled an "ecological inheritance" by Odling-Smee (1988).

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All organisms constantly interact with their local environments, and they
constantly change them by doing so. If, in each generation, populations of
organisms only modify their local environment idiosyncratically, or
inconsistently, then there will be no modification of natural selection
pressures, and hence, no significant evolutionary consequence. If, however,
in each generation, each organism repeatedly changes its own environment in
the same way, perhaps because each individual inherits the same genes
causing it to do so, then the result may be a modification of natural
selection.

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CB: Why do they say "perhaps" ? Isn't this definitely so. And isn't this an important 
point in a paper that compares instinctive behavior and cultural behavior ?

I can understand that the authors are pointing to ecological inheritance as structures 
external to the organisms's bodies and thereby different than the bodily phenotype. 
But the behaviors that generate the physical structures that persist to a next 
generation or influence the upbringing of infant animals here are INSTINCTIVE 
behaviors, not learned as a culture or tradition.

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Figure 1: (a) The standard evolutionary perspective: populations of
organisms transmit genes from one eneration to the next, under the direction
of natural selection. (b) With niche construction: phenotypes modify their
local environments (E) through niche construction. Each generation inherits
both genes and a legacy of modified selection pressures (ecological
inheritance) from ancestral organisms.

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CB: But the "modified selection pressures" do not modify anew each generation. They 
are the same for each new generation, because each previous generation "modifies" it 
in the same way.  The only modification would be between the environments of the first 
generation that aquired the instinctive behavior with a new gene  and the one before 
that didn't have the gene and instinctive behavior. 

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We have outlined the principal evolutionary consequences of niche
construction elsewhere (Odling-Smee et al., 1996; Laland et al., 1996a). Our
goal here is to spell out the repercussions of this perspective for the
human social sciences. We maintain that a focus on niche construction has
important implications for the relationship between genetic evolution and
cultural processes. The replacement of a single role for phenotypes in
evolution by a dual role immediately takes away from human culture its claim
to a unique status with respect to its capacity to modify natural selection.
Humans can and do modify many natural selection pressures in their
environments, but the same may be said of many species, and most do so
without the help of culture. Moreover, this dual role for phenotypes implies
that a complete understanding of the relationship between genes and culture
must acknowledge not only genetic and cultural inheritance, but also take
account of the legacy of modified selection pressures in environments. To
illustrate these points, we need to take a fresh look at how human culture
relates to human evolution in the light of niche construction.


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CB: Seems to me that human culture still has a unique status with respect to its 
capacity to modify natural selection. The examples the authors have given of 
"ecological inheritance" in non-human species are still modifications of the 
environment based in instinctive behavior.  Human culture creates non-instinctive 
behavior that not only modifies the environment and natural selection, but the 
behaviors themselves are passed on externally to the body, through symbols and 
education directly, not indirectly by the remaining modified niche.  There is still a 
qualitative difference in the way that human culture can modify natural selection. A 
beaver can leave behind a dam that will modify the environment of the next generation 
of beavers. But a beaver can't leave behind a blueprint or even better a description 
in words,  of how to build a dam to he next generation.  Marx's little ditty in 
_Capital_ on the difference between the construction by a spider or a bee is still a 
good rule of thumb on this. The human builder , unlike the spider or the bee, builds 
in imagination first, has a plan.  The enormity of the "imaginary plans" , culture, 
tradition, custom, that humans accumulate and pass on has gone from a quantitative 
difference to a qualitative difference with animal transgenerational transfers.   An 
animal can pass on an emblem ( a dam) , but only a human can pass on a symbolic 
description of how to build a dam. 


CB

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