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On Racial Memory

Racial Memory and Instinct: The Case of the Honeyguide

Like the Lernean Hydra, Lamarckism refuses to go away no matter how
many times it has been pronounced dead and buried by the leading
lights of science.  Having elsewhere discussed new research supporting
the possibility of directed mutation1i.e., mutations directed in
accordance with the needs of the evolving organism, the classic
Lamarckian positionwe will here review a recent field-study which
seems to suggest the necessity of some sort of racial memory.2

Across the bushlands of northern Kenya, a remarkable example of
cooperation between bird and man can be found, operating today much as
it has for untold centuries.3   This is the territory of the Boran, a
nomadic tribe who continue to subsist through hunting and gathering.
For as long as they can remember, the Boran have been taking advantage
of the helpful behavior of a bird known as the greater honeyguide
(Indicator indicator), whichas its name impliesis widely renowned as a
guide towards choice deposits of honey.

By nature the honeyguide displays a great fondness for beeswax and bee
larvae.  Inasmuch as beehives are virtually inaccessible to these
birds and well protected by the bees (Apis mellifira)  themselvesthe
colonies are typically located in large trees, rock crevices or
termite moundsthe honeyguide requires the aid of man in order to
pillage the honey-cache.  As the natives tell itand now confirmed by
the study of Isack and Reyerthey summon the honeyguide to the campsite
by emitting a specific, penetrating whistle which can be heard from a
distance of over one kilometer.4
Upon hearing the whistle, the bird responds by flying towards the
campsite of the Boran, whereupon it announces itself by flying close
to the humans and moving restlessly amongst conspicuous perches in the
trees, all the while emitting a very characteristic call
(tirr-tirr-tirr-tirr).  Once it has gained the attention of a
potential honey-gatherer, the bird then proceeds to guide him in a
fairly direct manner to the site of the bee-colony, the latter of
which may be several kilometers away.  Alternately appearing and
disappearing, the bird periodically returns to check on the humans, as
if to make sure that they are still following, such returns becoming
more frequent and agitated as the two parties near the hive.  The
Boran honey gatherers, in turn, whistle and bang sticks while
following the bird to maintain its attention and announce their
position.  According to professional honey gatherers, "through its
guiding pattern, the bird informs them about the direction of, the
distance to, and their arrival at the colony."5

Upon arriving at the site of the bees' nest, the honeyguide perches
near it and emits a very characteristic "indication" call.  By all
accounts, this call differs markedly from the guiding call.  Should
the Boran experience difficulty in locating the colony, the bird
swoops down and circles it as if to further clarify the precise
location of the nest.6  Once the nest is found and its thick walls
breached, the Boran plunder the honeycombs, always leaving a few
select morsels behind for the honeyguide.7

Although countless examples of cooperative behavior are known from the
annals of naturalists and nature lovers alike, the case of the
honeyguide is special because of the complexity of the behaviors and
interspecific communication involved, the latter apparently being the
product of multiple generations of development and refinement (in this
particular tete-a-tete, it would appear that man is a relatively
recent interloper, having replaced the ratelMellivora capensisthe
latter being a badger-like animal known to serve as juggernaut to the
honeyguide).8  Long considered the stuff of legend, the now
well-documented partnership pertaining between the greater honeyguide
and Boran nomads presents naturalists with a vexing problem: How is it
possible to explain the development and evolution of this particular
symbiotic relationship?

Certainly it seems highly unlikely that an ancestral honeyguide
suddenly appeared one day with a new mutation in its genome which
compelled it to seek out a friendly benefactor resilient enough to
conquer a colony of angry bees.  Fortuitous mutations of this
sortfrequently invoked in conventional interpretations with regard to
the origin of structural modificationsseem more than a little
far-fetched when attempting to account for the evolution of complex
behaviors.  Despite decades of experimentation, moreover, no mutation
has yet been observed to give rise to a new behavioral pattern, only
to pathological distortions of already existing behaviors.9

A more realistic approach would take its cue from the process known as
organic selection.10
Originally offered as a compromise between Lamarckism and Darwinian
selection, this theoryas its name impliesfollows Lamarck in supposing
that an organism's habitual behavior plays a decisive role in its
evolution.  Baldwin's summary of the idea is as follows:
"Acquired characters, or modifications, or individual
adaptationswhile not directly inherited, are yet influential in
determining the course of evolution indirectly.  For such
modifications and accommodations keep certain animals alive, in
this way screening the variations which they represent from the
action of natural selection and so allow new variations in the
same directions to arise in the next and following generations;
while variations in other directions are not thus kept alive and
so are lost.  The species will therefore make progress in the same
directions as those first marked out by the acquired modifications
and will gradually 'pick up' by congenital variation the same
characters which were at first only individually acquired.  The
result will be the same as to these characters, as if they had
been directly inherited."11

Confronted with the case of the honeyguide, organic selectionists
would presumably argue that the bird's behavioral complex originated
in the customary manner, through trial, error, and reinforcement over
the years.  As inveterate bee-watchers, it may be supposed that the
wax-loving honeyguides would have had plenty of opportunity to observe
ratels or humans in the act of plundering a bees' nest.  Upon
benefiting from scraps inadvertently left behind, the bird might well
have been tempted to follow its mammalian benefactor to see if future
feasts could be had.  If such an experience met with success, it is
understandable how a casual association between the two species could
have developed.12  Anyone who has ever shared table scraps with a
dogthe end result of which is that all future meals feature a
four-legged beggarwill readily understand how an association between
the mammal and honeyguide could have originated.

But how are to move from a casual relationship between two species to
one in which the bird chooses to act as a guide to the mammal in treks
across the African bush?  Supposing that honeyguides were already
accustomed to tagging along with ratels/humans during the search for
honey, it seems but a natural development for the bird to turn the
tables and lead the mammal to yet another bees' nest already known to
it, particularly if the latter nest lie in close proximity to one just
plundered.  Here all that need originally be involved was that the
bird direct the mammal's attention to a nearby nest, either by
circling it or by raising a ruckus.  No real guiding need be involved
in the incipient stages of the behavior.  In this way, perhaps, the
bird first gained the confidence of the ratel that it could help
locate honey.  Given further experience, practice, and the development
of mutual trust, it stands to reason that the bird might inspire the
mammal to pursue ever longer forays into the bush.  And before you
know it, a symbiotic relationship would develop to the point where
either party might approach the other, looking for assistance.13

While any number of "just so" scenarios are conceivable, it stands to
reason that the guiding behavior of the greater honeyguide originally
became established through learning and habituation in one
particularly clever or adventurous bird, thereafter spreadingeither
through imitation or communicationto its peers and progeny.14  So far
so good.  But how are we to move from a habit acquired in one
generation of birds to an instinctual behavior characteristic of the
species (that the behavior is indeed instinctive in nature is commonly
agreed upon15 and, in any case, is rendered near certain by the
unusual circumstance that honeyguides are parasitic breeders; i.e.,
like the cuckoo, they lay their eggs in the nests of other birds, and
thus the nestling(s) would appear to have little opportunity to learn
the intricacies of symbiotic honey-guiding from its parents)?16  Here
the organic selectionists would argue that if the symbiotic
honeyguides gained a competitive advantage over non-symbiotic birds in
their struggle for survival, this would favor the greater honeyguide,
giving it additional time during which the originally "acquired"
character (the bird's guiding behavior) would eventually be
supplemented by genetic factors, thereby becoming a permanent
characteristic of the species.

Yet it is far from clear how natural selection is to operate in the
present example.  Of the 17 species of honeyguides, only two17 are
known to participate in symbiotic honeyguiding, and thus it is
difficult to believe that such behavior is a matter of life and death
for the greater honeyguide (although it might well be granted that the
periodic raiding of a bee-colony produces certain nutritional benefits
as it entails certain risks).  Friedmann arrived at a similar
conclusion:
"The fact that many species of honey-guides do not guide (as far as
known) and yet get enough wax regularly (practically all specimens
collected have had wax in their gizzards) shows that guiding is
not necessary.  That it probably never was essential seems to be
reflected in the fact that it did not 'carry over' into the other
phylogenetic branches of the family."18

In short, if the greater honeyguide's propensity for guiding mammals
doesn't act so as to favor it over its wax-loving relatives in the
struggle for survival, there is nothing for selection to act upon and
thus it can play no role in the genetic stabilization of the
behavior.  And if the operation of selection is difficult to imagine
when the guiding behavior is fully developed, how is it to act during
the presumably lengthy period when the behavior was first appearing
and developing by incremental stages?  Here Friedmann remarked: "It
is, however, difficult if not impossible to imagine the development of
such a habit if it were the chief foraging method, as it would have
been of no conceivable value to either until it was perfected by
both."19

As critics have pointed out, a far more serious problem faces
proponents of organic selection in trying to account for the
transition of the bird's behavior from an acquired trait to a genetic
one: If greater honeyguides can survive the rigors of existence
through learned symbiotic behavior alone, how is it that selection
enters the picture to insure that random mutations accumulate in the
right direction in order to render the behavior instinctual in
nature?  Simply statedIt can't.20  And in the absence of selection,
proponents of organic selection can only say a little prayer and
suppose that the honeyguide's behavior ultimately becomes established
through a series of especially fortuitous mutations in the bird's
genome.

Questions arise at every step of this explanation.  For how many
generations must the honeyguide wait around for random genetic errors
to stabilize its behavior so it won't have to go through the
bothersome and inherently risky process of teaching its progeny such
sophisticated strategies for acquiring beeswax (here supposing, for
the sake of argument and in contradiction to the published reports,
that parental education plays a role in the symbiotic behavior pursued
by the honeyguide)?  And how many mutations would be required to
establish this remarkable behavioral complex?  Certainly it would be
asking a lot of one fortuitous error in genetic replication to encode
the honeyguide's various calls, object-oriented guiding, luring and
signaling behavior, knowledge of bee-behavior, etc.  Yet with each
additional fortuitous mutation invoked to account for the behavior's
respective components the problems are compounded.  One is reminded of
Koestler's description of Neo-Darwinian evolution as comparable to a
monkey pounding away randomly at the typewriter in the hopes that a
Shakespearean sonnet will result.21

But even granting that a genetic mutation occasionally proves
beneficial, how are we to account for the genome's apparent
acquisition of information about the environment?  For in the present
case it stands to reason that for the greater honeyguide's behavior to
have a genetic basis, its genome must have somehow acquired direct and
highly specific information about the goings-on in the Kenyan
bushlands.  At the very least one must suppose that the honeyguide is
born with an innate "knowledge" of bees, ratels, humans, beeswax,
guiding tactics, specific calls, etc.

Granted that knowledge is necessary to account for the honeyguide's
behavioral repertoirenow become instinctualthrough what mysterious
process does an avian genome acquire such knowledge?  Konrad Lorenz,
arguably the world's foremost authority on instinctual behavior,
offered the following explanation: "The adaptation of the behavior
patterns of an organism to its environment is achieved in exactly the
same manner as that of its organs, that is to say on the basis of
information which the species has gained in the course of its
evolution by the age-old method of mutation and selection."22  Lorenz'
stature as an ethologist notwithstanding, his statement is thoroughly
misleading as it is quite impossible that a mutation could provide an
organism with information about the environmentin the present case,
information regarding the intricacies of guiding behavior to a bird
hitherto ignorant of such practices.23

We thus seem to have reached an impasse in our quest to understand the
behavior of the greater honeyguide.  To suppose that a honeyguide can
perform the manifold behaviors attributed to it without any prior
experience, learning, or knowledge being required would appear a most
untenable position from the standpoint of animal psychology.  To
suppose, on the other hand, that a random error(s) in genetic
replication could somehow provide an untrained bird with the very
knowledge necessary for pursuing its remarkable symbiotic relationship
with the Boran nomads is tantamount to invoking a miracle.

But if the Neo-Darwinian theory of evolution is incapable of
explaining the behavior of the honeyguideand it iswhat is the
alternative to a blind faith in miracles, whether of the Neo-Darwinian
or Creationist variety?  If we might be allowed to entertain the
Lamarckian heresy, an obvious solution to the case of the honeyguide
presents itselfnamely, racial memory.  For Lamarck and Neo-Lamarckians
alike, instinct is to be understood as racial memory or habit,
behavior acquired as a result of the trials and travails of experience
and subsequently incorporated into the genome by some admittedly
mysterious process.  According to this theory, the individual
honeyguide behaves as it does not from personal experience or parental
education, rather as the beneficiary of the cumulative experience and
knowledge acquired by generations of ancestors, for whom the symbiotic
interaction with mammals was a long-established habit.

In order to understand the simplicity, logical force, and breadth of
appeal of the Lamarckian position, it is necessary to briefly review
its history.

Racial Memory: Lamarck and Darwin

Jean-Baptiste Pierre-Antoine de Lamarck is rightly regarded as the
originator of the first systematic theory of evolution.  Lamarck's
magnum opus, Philosophie zoologique, was published in 1809 and
subsequently languished in obscurity until Darwin's On the Origin of
Species made evolution a household word and the cornerstone of the
biological sciences (the word biology was actually coined by
Lamarck).

In Lamarck's system, an animal's behavior played a decisive role in
its evolution, both in selecting its environment and in shaping
structure through habitual function.  According to Lamarck, an animal
developed habitual patterns of behavior in response to various
environmental contingencies and in an effort to accommodate felt
needs.  Such habitual patterns of behavior, in turn, led to changes in
function and form.  Habitual behavior, combined with heredity, gave
rise to instincts:
"As the penchants that animals have acquired through the habits they
have been forced to contract have little by little modified their
internal organizations, thus rendering the exercise of (these
penchants) very easy, the modifications acquired in the
organization of each race are then propagated to new individuals
through generation.  It results from this that the same penchants
already exist in the new individuals of each race even before they
have the chance to exercise them."24

Lamarck's explanation of instinct as racial memory, although it has
forbears, actually represented a significant breakthrough inasmuch as
it was commonly held at the time that animals' instincts were the
product of special creation (design by God).25  And as the first
scientific explanation of instinct offered from an evolutionary
perspective, Lamarck's theory was destined to have a decisive
influence upon subsequent discussions in the field.  With the notable
exceptions of August Weismann and Alfred Wallace, nearly every
prominent naturalist of the nineteenth century adopted a "Lamarckian"
explanation of instinct, including Darwin, Romanes, and Haeckel.26

Darwin himself, as is well-documented, advocated the Lamarckian view
of instinct as racial memory in no uncertain terms.  Thus in an early
notebook we read: "Now if the memory of a tune and words can thus lie
dormant during a whole lifetime, quite unconsciouslysurely memory from
one generation to another also without consciousness, as instincts
are, is not so very wonderful."27  In the pioneering work The
Expression of the Emotions in Man and Animals, Darwin wrote of
instincts that "some cannot be distinguished from inherited
habits."28  Elsewhere we find Darwin writing as follows: "Some
intelligent actions, after being performed during several generations,
become converted into instincts and are inherited, as when birds on
oceanic islands learn to avoid man."29

The idea that innate behavior and instinct depend on racial memory and
habit pervades Darwin's discussions of emotion, sexual selection, the
origin of articulate speech, and the inordinate development of the
human brain.30  Indeed, it is impossible to understand Darwin's
writings in these areas apart from this principle.  It was the
difficulty of explaining the development of the human brain through
selection alone, it will be remembered, which had so bedeviled Wallace
that he sought recourse in mysticism.31  Unlike Wallace, Darwin had
Lamarckian inheritance to fall back upon in order to explain the
anomaly represented by the human brain.  For Darwin, the evolution of
the human brain was closely correlated with the origin of language,
the latter arising through the inherited effects of use of the vocal
chords in our primitive grunting ancestors.  Comparing our vocal
chords with those of the apes, Darwin observed that: "In man the
closely similar vocal organs have become adapted through the inherited
effects of use for the utterance of articulate language."32  Although
the development of language was of obvious selective value, Darwin
adds that "the relation between the continued use of language and the
development of the brain, has no doubt been far more important."33  In
this way, Darwin accounted for the inordinate development of the human
brain: "As the various mental faculties developed themselves the brain
would almost certainly become larger."34

I, for one, find it very difficult to find fault with this argument.35

Herbert Spencer and the Foundation of Evolutionary Psychology

Herbert Spencer, a largely self-educated man whose life spanned the
death of both Lamarck and Darwin, has been called the most influential
thinker of the nineteenth-century (Spencer was born in 1820, and died
in 1903).36  His contributions to evolutionary theory were manifold,
although they are seldom acknowledged by historians of biological
thought.37  It was Spencer who first used the word evolution in its
modern sense, signifying a change in species over time and involving
an increasing differentiation of structure and specialization of
function, giving rise to successively higher levels of integration (In
the first five editions of the Origin, Darwin never once used the word
"evolution").38 And it was Spencer who first offered evolutionism as a
unified philosophy of nature.39  Robert Carneiro offered the following
assessment of Spencer's contributions in this area:
"Not only did Spencer anticipate many of the ideas of modern
evolutionism, he also expressed them more clearly, more precisely,
and more forcefully than almost anyone has since.  We still have
much to learn from Herbert Spencer"40

Although it is fashionable nowadays to credit Darwin with laying the
foundations for modern psychology, Darwin himself acknowledged the
priority of Spencer.  Looking to the future, Darwin wrote: "Psychology
will be securely based on the foundation already well laid by Mr.
Herbert Spencer."41

Spencer's primary contribution to the field was the book The
Principles of Psychology (1855), whichalthough it drew little acclaim
at the time it was publishedwas destined to have a seminal influence
upon future psychologies.42  Spencer revolutionized the study of
behavior through his attempt to interpret it in terms of evolution.
Lamarck's theory of evolution, together with the mechanism of the
inheritance of acquired characters, served as the twin cornerstones of
Spencer's psychology.
His view that evolutionary history was the single most important
factor in the study of mental phenomena can be glimpsed in the
following quote: "If the doctrine of evolution is true, the inevitable
implication is that Mind can be understood only by observing how Mind
is evolved."43

An associationist in the tradition of Bain, Spencer was among the
first to attempt to understand Mind as a product of interaction with
shifting environments.  Thus he is rightly regarded as the founder of
the psychology of adaptation.44  Spencer was also among the first to
emphasize the importance of heredity.  As Hoffding noted, prior to his
time heredity had been regarded as little more than a curiosity, but
from Spencer onward "it was regarded as an ever-present cooperative
factor in the determination of the very highest forms of life."45

Spencer's evolutionary perspective, together with the concept of
Lamarckian inheritance, allowed him to offer a solution to the
hitherto intractable debate between the empiricists (Locke, Mill)who
held that all mental phenomena arose from experience, and thus no a
priori or innate ideas were possibleand the Kantians, who held that
innate mental categories organized experiences into coherent
relations.  Here Spencer argued that there was substantial truth in
both camps: All mental operations do stem from experience, but there
are also innate categories of mental relations, the all-important
catch being that the latter arise from the inherited effects of
evolutionary experience.46  In The Principles of Psychology, Spencer
argued as follows:
"Let it be granted that in all creatures, as in ourselves, the law is
and has been, that the more frequently psychical states occur in a
certain order, the stronger becomes their tendency to cohere in
that order, until they at last become inseparable; let it be
granted that this tendency is, in however slight a degree,
inherited, so that if the experiences remain the same, each
successive generation bequeaths a somewhat increased tendency; and
it follows, thatthere must inevitably be established an automatic
connection of nervous actions, corresponding to the external
relations perpetually experienced."47

Instinctual behavior had long represented a thorn in the side of
empiricists.  How could a bird's ability to build a nest be the
product of experience or education when a bird raised in isolation
likewise fashioned perfectly adequate nests?  The answer, according to
Spencer, was Lamarckian inheritance.  Instincts are simply associative
habits so well learned that they have become part of the genetic
heritage of the species: "The doctrine that the connections among our
ideas are determined by experience must, in consistency, be extended
not only to all the connections established by the accumulated
experiences of every individual but all those established by the
accumulated experiences of every race."48

Near the end of his career, Spencer became embroiled in a memorable
debate with August Weismann, the latter being the leading proponent of
the position that would come to be known as Neo-Darwinism (i.e, the
view that all evolutionary change results from natural selection
acting upon random changes in the genome, thereby excluding any and
all Lamarckian effects).49  It was Weismann's arguments against the
possibility of the inheritance of acquired characters that began to
turn the tide against Lamarckian views in the last two decades of the
nineteenth century.50
Spencer's rejoinder to Weismann stands as one of the classic
defenses of Lamarckian evolution.  In addition to presenting evidence
favoring the Lamarckian position (the blindness of many cave animals,
an apparent example of the degeneration of structure through disuse;
the inordinate sensitivity of the human tongue, attributable to the
inherited effects of manipulating food and rubbing up against teeth;
etc.), Spencer also offered various challenges to the efficacy of
natural selection as an agent of evolutionary change.51  There Spencer
offered his famous statement: "Close contemplation of the facts
impresses me more strongly than ever with the two alternativeseither
there has been the inheritance of acquired characters, or there has
been no evolution."52

It would be impossible here to provide an adequate assessment of the
arguments of Spencer and Weismann.  Suffice it to observe that modern
biology has tended to follow Weismann's view of the evolutionary
process.  It is not without interest, however, that Weismann would
abandon his faith in the "all-sufficiency" of natural selection
shortly thereafter, in large part because of Spencer's critique.
Modern evolutionary theory, with its belief in the "all-sufficiency"
of natural selection and concomitant rejection of Lamarckian
inheritance, thus stands in the curious position of advocating a
theory abandoned by Weismann and emphatically disavowed by Darwin
himself.

In his autobiography, while contemplating the ever-growing influence
of Weismannism in biological circles, Spencer would declare: "I still
hold, as I then held, that the inheritance of functionally wrought
modifications is the chief and most exclusive factor in the genesis of
all the more complex instincts and all the higher mental powers."53

Darwin and Spencer were hardly alone in their defense of the
Lamarckian view of instinct as racial memory.  According to Mayr, many
naturalists continued to believe in Lamarckian inheritance until
relatively recently (c. 1930-1950).54

But what of the various schools of psychology whose business it is to
investigate and analyze the phenomena of animal behavior and
instinct?  As I have documented elsewhere55, a survey of pivotal
figures in the history of psychologyWundt, Hall, Freud, Jung, Watson,
Pavlov, Macdougall, Bleulerwill reveal that most were indefatigable
proponents of the doctrine of the inheritance of acquired characters,
this despite the fact that it would be difficult to find another issue
upon which these men would agree.56

At this point a brief review of the opinions of various prominent
psychologists is in order.

Racial Memory in the Annals of Psychology

Wilhelm Wundt is rightly regarded as a founding father of experimental
psychology, but his influence upon physiological, social, and
developmental psychology was also considerable.57  Long occupied with
the phenomena of instinct, Wundt believed that instincts were
originally volitionalsometimes consciousbehaviors gained, practiced,
and improved upon through the generations.  Drawing an analogy between
complex instincts such as a bird's ability to build a nest and the
ability to play a piano, Wundt noted that even highly complex
behaviors can become nearly automatic with sufficient practice:
"We may, accordingly, explain the complex instincts as developed
forms of originally simple impulses which have gradually
differentiated more and more in the course of numberless
generations, through the gradual accumulation of habits which have
been acquired by individuals then transmittedThe gradual passage
of a connate disposition is to be explained as a psycho-physical
process of practice through which complex volitional acts
gradually pass into automatic movements following immediately and
reflexly the appropriate impression."58

Like Wundt, Sigmund Freud remained an ardent Lamarckian throughout his
long life.  Without this element, much of what Freud says on
individual and group psychology, anthropology, myth, symbolism,
religion, and the aetiology of neuroses would not be fully
comprehensible.59

For Freud, instinct was racial memory pure and simple, and mankind's
racial heritage was no different in this respect than that of any
other living creature:
"[The concept of ancestral memories helps] diminish the overwide
gap human arrogance in former times created between man and
beast.  If the so-called instincts of animalswhich from the very
beginning allow them to behave in their new conditions of living
as if they were old and long established onesif this instinctual
life of animals permits of any explanation at all, it can only be
this: that they carry over into their new existence the experience
of their kind; that is to say, that they have preserved in their
minds memories of what their ancestors experienced.  In the human
animal things should not be fundamentally different.  His own
archaic heritage, though different in extent and character,
corresponds to the instincts of animals."60

Fully cognizant that Lamarckism was rapidly losing favor with his
colleagues in biology, Freud reiterated his position on the great
heresy in his last book: "I admit, in all modesty, that in spite of
this I cannot picture biological development proceeding without taking
this factor into account."61

William McDougall made a name for himself in the first four decades of
this century as a pioneering social psychologist who stressed the
instinctual basis of human behavior and as an outspoken critic of
Watson's behaviorism.  Indeed, his Social Psychology (1907) is still
regarded as a "founding book" in the field.62  McDougall emphasized
the purposive aspects of behavior and instinct, arguing that their
evolution demanded a Lamarckian explanation.  Indeed, McDougall went
so far as to argue that without the Lamarckian principle, "our
understanding of man's nature, of organic evolution, and of historical
development, all alike are at a deadend."63

McDougall subsequently gained a fair amount of notoriety when he
announced positive results in an extensive experiment designed to test
the heritability of the effects of training in rats.64  Described by
an historian of science as "one of the most elaborate biological
experiments yet devised,"65 McDougall's experiment spanned several
decades.  Near the end of his career, McDougall underscored the
importance of the debate over Lamarckian inheritance:
"The question in dispute not only lies at the very heart of the
problem of evolutionbut also it is at the heart of the greater
issue between, on the one hand, scientific materialism and, on the
other hand, every form of science, philosophy, and religion that
would regard mind as in any sense real and operative in the
universe."66

Summary

Modern evolutionary theory, it has been argued, is completely
incapable of explaining the development of the unique symbiosis
pertaining between the greater honeyguide and various mammals.  We
have seen that it is difficult to imagine how selection is to operate
with regards to the fully developed symbiosis, much less during the
incipient stages of the behavioral complex throughout that presumably
long period when it would have been of little or no use to either bird
or mammal.  The Lamarckian theory, on the other hand, explains the
symbiosis with relative ease and in a common sense manner which every
layman can appreciate.  The greater honeyguide acts as it does because
its ancestors acquired extensive knowledge of mammals and bees during
their long history in the African bush, such knowledge subsequently
being transmitted to progeny through the inheritance of acquired
characters.  No experience or practice is necessary for the individual
honeyguide to perform its dazzling array of behaviors for the simple
reason that its ancestors have already experienced, practiced, and
memorized the intricacies of honey-gathering through the mediation of
mammals.  The Lamarckian hypothesis has as an additional point in its
favor the fact that it alone can explain the development of the
behavioral complex during its incipient stages, prior to it serving
any utilitarian function.  All that is necessary is that associations
in memory develop between the honeyguide and mammal and that these
learned associations are heritable in some degree.  If such be
granted, it is possible to understand how an originally fortuitous
meeting of a honeyguide and ratel at a beehive couldgiven numerous
such meetings and several generationsdevelop to the point where
additional associations, mutual trust, the requisite modes of
communication, etc., would further refine the habitual operation of
the symbiosis.

To this simple explanation of the honeyguide's behavior it will no
doubt be objected that there is as yet no known biochemical mechanism
whereby characters acquired by the individual organism might be
transmitted to its genome.  The following points are relevant here:
(1) the history of science bears witness that there is ample precedent
for setting aside conceptual difficulties when the evidence or logic
bearing upon a particular matter warrants it; and (2) recent
developments in genetics make the possibility of Lamarckian
inheritance increasingly likely.

With regards to the first point, there is the example provided by
Newton's theory of gravitation, long considered the very ideal of
scientific reasoning.  We all take the reality of gravity for granted,
but who can explain the physical mechanism behind its operation?  Sir
Isaac himself didn't pretend to know the answer:
"It is inconceivable, that inanimate brute matter should, without the
mediation of something else, which is not material, operate upon,
and affect other matter without mutual contactAnd this is one
reason, why I desired you would not ascribe innate gravity to me.
That gravity should be innate, inherent, and essential to matter,
so that one body may act upon another, at a distance through a
vacuum, without the mediation of anything else, by and through
which their action and force may be conveyed from one to another,
is to me so great an absurdity, that I believe no man who has in
philosophical matters a competent faculty of thinking, can ever
fall into it.  Gravity must be caused by an agent acting
constantly according to certain laws; but whether this agent be
material or immaterial, I have left to the consideration of my
readers."67

The evidence supporting racial memory, if not as conclusive as that
supporting Newton's theory, is nevertheless compelling.  As to the
actual biochemical mechanism(s) behind the phenomenon of racial
memory, I confess to being as much in the dark about it as about the
mechanism behind gravity itself.  Future discoveries, I trust, will
enlighten us on both fronts.

With regards to recent developments in genetics, signs abound that the
Neo-Darwinian conception of the genome as an isolated system
impervious to external influences and ruled only by random errors in
replication is nearing extinction.  Consider the following observation
offered by a distinguished geneticist: "Genomes function as true
intelligent systems which can be readjusted when conditions
require."68  Here is a  statement all Lamarckians could endorse.

That accidents in genetic replication occasionally occur goes without
saying.  That on very rare occasions, such accidents subsequently
prove beneficial to the evolving organism is possible, perhaps even
likely.  But it by no means follows that such inherently accidental
phenomena govern the process of genetic variation and, consequently,
adaptation and evolution as well.

Certainly it is no accident that the genome has evolved highly
efficient and sophisticated strategies for checking and correcting the
inevitable errors in replication.  And however we choose to label such
self-regulating processes, there would seem to be little doubt but
that the genome is constantly monitoring its various components and
products in accordance with some "model" of self, health, and
adaptation.  Are we to suppose, then, that the genome has an inherent
ability to monitor and correct its own products but no power to direct
new mutations in order to accommodate the ever-changing needs of the
cell/organism?

At the very least, these well attested systems of auto-regulation
confirm that the genome is continually receiving, processing, and
organizing information about its environment.  How such organization
takes place, we have very little idea.  How extensive is the nature
and informational content of the feedback involved is anybody's
guess.  All that we know for certain is that some such feedback must
be involved if we are to make sense of the phenomena associated with
the evolution of the genome, cellular differentiation, and
regeneration.

Nor should it be forgotten that the genome itself is composed of
originally separate organisms which, once-upon-a-time, elected to
commingle and cooperate in order to insure the survival of the
respective selves.69  That such cooperation necessarily requires
incessant communication, feedback, and genetic fine-tuning between the
respective parties is both obvious and undeniable.  Indeed, it is
arguable that the interspecific communication which pertains between
the various endosymbionts of the genome is functionally analogous with
that observed to operate between the greater honeyguide and the Boran
nomad.  Both originated as a result of evolutionary experience and
biological need which, given practice and refinement, eventually
became habitual and genetically conditioned.  And both are decidedly
goal-directed and thus purposive in nature.

It is my opinion that the many wonders of instinct, adaptation, and
evolution can best be understood if genetic change is an ordered
phenomenon; i.e., subject to auto-regulation on the part of the
evolving genome/organism.  Genomic evolution is simply the
bio-chemical manifestation of an organism's acquisition of information
about its environment, learning on the grand scale of phylogeny as
opposed to ontogeny.70  As the organism comes to learn more about its
environment, its genome responds with the appropriate changes in order
to insure that the "knowledge" gained is preserved for posterity.  As
mysterious as the biological processes which would allow for such
informational feedback between the organism and its genome currently
seem, they are no more mysterious than those processes which allow for
cellular differentiationhere, it will be remembered, each of the
functionally different cells has the same store of genetic
information, so who or what is directing the epigenetic process and by
what biochemical means?  How, apart from some form of detailed
feedback between the developing "somatic" cells and the genome is it
possible to account for the facts of cellular differentiation?  Yet
once grant the possibility of such feedback between somatic cells and
the genome during embryogenesis and it becomes impossible to deny
similar means of feedback during the life of the organism.

The case of the honeyguide presents biologists with a dilemma: Do we
accept the perfectly logical Lamarckian explanation of the symbiosis
between the greater honeyguide and mammals as reflecting the inherited
effects of experience and habit even though we do not yet know of any
biochemical mechanism whereby such inheritance might take place?  Or
do we accept the Neo-Darwinian explanation of the symbiosis as a
product of random mutation aided by natural selection, even though it
is difficult if not impossible to imagine how selection might be
involved?  If racial memories are heritable, the honeyguide's
instinctual behavior is rendered perfectly understandable.  Only the
precise biochemical means of the encoding and transmission of such
racial memories remains mysterious.  If, on the other hand, there is
no such thing as racial memory, the case of the honeyguidelike a
myriad of similar examples from the world of animalsmust remain
inexplicable and more than a little miraculous.

Footnotes

1. E. Cochrane, "Viva Lamarck: Renewed Discussion on the Inheritance
of Acquired Characters," Aeon 2:2 (1991), p. 5-39.  See also the
valuable discussion in E. Fox Keller, "Between Language and Science:
The Question of Directed Mutation in Molecular Genetics," Perspectives
in Biology and Medicine (1992), pp. 292-306.

2. The definitive study of this bird is that of H. Friedmann, "The
Honey-guides," U.S. Nat. Mus. Bull. 208 (1955), pp. 1-292.  According
to Friedmann, reports describing the guiding behavior of the
honeyguide are attested as early as 1569.

3. H. Isack & H. Reyer, "Honeyguides and Honey Gatherers:
Interspecific Communication in a Symbiotic Relationship," Science 243
(1989), p. 1344.  There the authors present the results of a
three-year study in which they documented to a great extent the
seemingly far-fetched claims of the Boran honey-gatherers that these
birds use a wide variety of very specific behaviors to lead them to a
bee-hive.  D. Griffin, Animal Minds (Chicago, 1992), pp. 164-169
provides an overview of the findings of Isack and Reyer and compares
them with earlier accounts.

4. Ibid., p. 1344.

5. Ibid., p. 1345.

6. Ibid., pp. 1343-1346.

7. R. May, "Honeyguides and Humans," Nature 338 (1989), p. 707.  See
also the discusion of H. Friedmann, op. cit., pp. 41-50.

8. R. Wesson, Beyond Natural Selection (Cambridge, 1993), p. 241.
There Wesson quotes Fuller and Thompson to the following effect: "No
gene has been demonstrated to direct a complicated series of motor
responses that produce an integrated behavior pattern."

9. The same general concept appears under various other names as well,
including the Baldwin effect (after the famous psychologist James Mark
Baldwin) and genetic assimilation (a phrase coined by C.H.
Waddington).  Originally proposed by Baldwin, Morgan, and Cope in the
past century, this theory has been defended most recently by Sir
Alistair Hardy, The Living Stream (New York, 1965).  See the extensive
discussion in Ev Cochrane, "Psychology, psychologists, and evolution,"
(Ames, 1981), pp. 127-160.  [Master's thesis submitted to Iowa State
University]  See also R. Richards, Darwin and the Emergence of
Evolutionary Theories of Behavior (Chicago, 1987), pp. 451-503; G.
Simpson, "The Baldwin Effect," Evolution 7 (1953), pp. 110-117; C.
Waddington's "The evolution of adaptations," Endeavor (July, 1953),
pp. 134-139.

10. J. Baldwin, Development and Evolution (New York, 1902), pp.
137-138.

11. In a similar manner, birds typically tag along with large
herbivores to feast upon the insects that hover about the mammals.

12. For a similar attempt at reconstructing the origins of the
honeyguide's behavior, see H. Friedmann, op. cit., pp. 66-71.

13. According to H. Friedmann, op. cit., pp. 33, 67, honeyguides are
typically solitary birds and thus it is that the vast majority of
guiding excursions are led by one bird.

14. H. Friedmann, op. cit., p. 54, calls it a "purely instinctive
behavior pattern."

15. L. Short & J. Horne, "Behavioral Notes on the Nest-Parasitic
Afrotropical Honeyguides (Aves: Indicatoridae)," American Museum
Novitates 2825 (1985), pp. 40-42.   Inasmuch as honeyguides appear to
monitor the progress of the nestling's infancy from afar, the
possibility exists that vocal communication could play some role in
their education into the ways of honeyguiding.  That a "culture" of
honeyguiding exists among this species, thereby accounting for the
behavior in individual birds, seems highly unlikely.

16. In addition to the greater honeyguide, Indicator variegatus is
also known to guide.

17. H. Friedmann, op. cit., pp. 65-66.

18. H. Friedmann., op. cit., p. 70.

19. See here the discussion of R. Richards, op. cit., pp. 494-495.
There he writes as follows: "If individual accommodations were
sufficient to preserve an organism, then congenital variations would
have little or no utility and thus would not be naturally selected
for."

20. In The Case of the Midwife Toad (New York, 1972), p. 30, Arthur
Koestler wrote as follows: "Neo-Darwinism does indeed carry the
nineteenth-century brand of materialism to its extreme limitsto the
proverbial monkey at the typewriter, hitting by pure chance on the
proper keys to produce a Shakespeare sonnet."  In a review of
Koestler's book, Stephen Gould offered the following retort: "The
simile is hoary enough, but it is utterly inappropriate.  Rather, our
monkeys must be allowed to keep all the correct letters after each
trial.  'Fortune and men's eyes' will soon arise."  "Zealous
Advocates," Science 176 (May 12, 1972), p. 624.  In the current
example, as in many others, Gould's objection is entirely unfounded,
there being nothing for selection to act upon.  Nor, as we will see,
is there any way for random mutations to provide a genome with the
necessary experience/knowledge to explain the bird's behavior.

21. K. Lorenz, in the preface to Darwin's The Expression of the
Emotions in Man and Animals (Chicago, 1965), p. xiii.

22. Properly speaking, a mutation can never cause an organism to
acquire information about its environment, for such would require that
the environment already pre-existed in some sense in its genome.
Indeed, there would appear to be but one way for a genome to acquire
new information according to orthodox genetic theory, and that is by
the incorporation of extraneous genetic elements such as episomes and
other factors.

23. Quoted from R. Burkhardt, The Spirit of System: Lamarck and
Evolutionary Biology (Cambridge, 1977), p. 170.

24. For a thorough review of ideas regarding instinct in the 17th and
18th centuries, see R. Richards, Darwin and the Emergence of
Evolutionary Theories of Behavior (Chicago, 1987), pp. 22-31.

25. Darwin, it is true, alternately defended the Lamarckian
explanation of instinct as racial habit and the "Darwinian"
explanation through selection alone.  Darwin appears to have been led
to this uncomfortable position as a result of his inability to explain
the complex instincts of the social insects through the Lamarckian
position.  See here the discussion of R. Richards, Darwin and the
Emergence of Evolutionary Theories of Mind and Behavior (Chicago,
1987), pp. 142-152.  For a Lamarckian explanation of the instincts of
social insects see Samuel Butler, Life and Habit (London, 1981), pp.
220-252; H. Spencer, Synthetic Philosophy (London, 1898), pp. 662ff.
I plan to return to this issue in a future essay.

26. C. Darwin, Metaphysics, Materialism & the Evolution of Mind
(Chicago, 1980), p. 7.

27. C. Darwin, The Expression of the Emotions in Man and Animals
(Chicago, 1965), p. 35.

28. C. Darwin, The Descent of Man (New York, 1874), p. 76.

29. See the discussion in E. Cochrane, "Psychology, psychologists, and
evolution," (Ames, 1981), pp. 22-30.

30. A. Wallace, Natural Selection (London, 1870).  On this question,
see also the valuable discussion of R. Richards, op. cit., pp.
176-184; and S. Gould, The Panda's Thumb (New York, 1980), pp. 47-58.

31. C. Darwin, The Descent of Man (New York, 1874), p. 56.

32. Ibid., p. 99.

33. Ibid., p. 60.

34. Interestingly enough, Darwin traced the religious impulseat least
in partto Lamarckian effects.  Of mankind's tendency to believe in a
First Cause (or a god), he wrote that it "probably depends merely on
inherited experience," adding: "Nor must we overlook the probability
of the constant inculcation in a belief in God on the minds of
children producing so strong and perhaps an inherited effect on their
brains not yet fully developed, that it would be as difficult for them
to throw off their belief in God as for a monkey to throw off its
instinctive fear and hatred of a snake."  See N. Barlow ed., The
Autobiography of Charles Darwin (New York, 1958), p. 93.

35. R. Hofstadter, Social Darwinism in American Thought (Boston,
1955).  See also the discussion of R. Young, Mind, Brain and
Adaptation in the Nineteenth Century (Oxford, 1970), pp. 190-196.   In
addition to documenting the immense influence of Spencer's psychology,
Young gives a very thorough summary of the development of his thought.

36. The worst example of this tendency to delegate Spencer to the
dustbins of history is that of Mayr, who wrote that "It would be quite
justifiable to ignore Spencer totally in a history of biological ideas
because his positive contributions were nil."  See E. Mayr, The Growth
of Biological Thought (Cambridge, 1982), p. 386.  More sober
assessments of Spencer's influence have been rendered by R. Roberts,
Darwin and the Emergence of Evolutionary Theories of Mind and Behavior
(Chicago, 1987), pp. 243-330; and M. Ridley, "Coadaptation and the
Inadequacy of Natural Selection," British Journal for the History of
Science 15 (1982), pp. 45-68.

37. See the comments of R. Carneiro in D. Freeman, "The Evolutionary
Theories of Charles Darwin and Herbert Spencer," Current Anthropology
15:3 (1974), p. 223.   According to P. Bowler, "Changing Meaning of
'Evolution'," Journal of the History of Ideas 36 (1975), p. 111, the
word evolution came into common use with the popularity of Spencer's
philosophy.

38. As Hoffding has pointed out, Spencer's vast philosophical system
can be regarded as the application of Lamarckian evolutionary
principles to the world and science at large.  H. Hoffding, A History
of Modern Philosophy (New York, 1955).  C. Smith, similarly, has
written that: "Spencer may be regarded as the nineteenth century's
philosopher of evolution.  His entire synthesis is pervaded by the
idea."  C. U. Smith, "Evolution and Mind," J. of the History of
Biology 15:1 (1982), p. 57.

39. R. L. Carneiro, "Structure, function, and equilibrium in the
evolutionism of Herbert Spencer," J. of Anthropological Research 29
(1973), p. 94.

40. C. Darwin, On the Origin of Species (New York, 1928), pp. 461-462.

41. It is known that Spencer's formulations had a profound effect upon
the thinking of Georges Romanes, a leading pioneer in the study of
animal behavior; upon William James, the founder of functional
psychology; upon James Watson, the originator of behaviorism; and upon
Sigmund Freud, the father of psychoanalysis.  See here the extensive
discussion of Robert Young, Mind, Brain and Adaptation in the 19th
Century (Oxford, 1970), pp. 191-196.  On Spencer's influencethrough
John Hughlings Jacksonon Freud, see F. Sulloway, Freud, Biologist of
the Mind (New York, 1979), p. 270ff.

42. H. Spencer, Principles
of Psychology Vol. 1 (London, 1870), p. 291.

43. T. Leahey, A History of Psychology (Englewood Cliffs, 1980), p.
246.

44. H. Hoffding, op. cit., p. 455.

45. Here, too, Darwin endorsed the opinion of Spencer.  In The Descent
of Man, p. 111, for example, Darwin wrote: "The ignoring of all
transmitted mental qualities will, as it seems to me, be hereafter
judged as a most serious defect in the works of Mr. Mill."

46. H. Spencer, Principles
of Psychology Vol. 1 (London, 1870), pp. 547-548.  See here the
discussion of J. Greene, "Darwin as a Social Evolutionist," Journal of
the History of Biology 10:1 (1977), p. 6.

47. H. Spencer, op. cit., p. 529.

48. H. Spencer, "The Inadequacy of Natural Selection," Contemporary
Review 43 (1893), pp. 153-166, 439-456; "Professor Weismann's
Theories," Contemporary Review 43 (1893), pp. 743-760.

49. In 1889, George Romanesalthough himself an ardent Lamarckiancould
speak of the widespread abandonment of Lamarckism as a result of
Weismann's ideas.  G. Romanes, An Examination of Weismannism (Chicago,
1893).  See also the discussion in D. Freeman, "The Evolutionary
Theories of Charles Darwin and Herbert Spencer," Current Anthropology
15:3 (1974), p. 216.

50. Most important here, perhaps, is the argument that natural
selection cannot account for the evolution of complex coadaptations.
See the discussion of M. Ridley, "Coadaptation and the Inadequacy of
Natural Selection," British Journal for the History of Science 15
(1982), pp. 45-68.  Ridley, p. 48, quotes Sewall Wright as follows:
"The apparent inadequacy of Darwin's theory to account for the
extraordinary coadaptive nature of organisms and the frequent apparent
uselessness of the components of coadaptive systems by themselveswas
probably the main reason that why most biologists, even into the 20th
century, continued to accept the inheritance of acquired characters."

51. H. Spencer, "The Inadequacy of Natural Selection," Contemporary
Review 43 (1893), p. 446.

52. D. Duncan, The Life and Letters of Herbert Spencer (London, 1911),
p. 547.

53. E. Mayr & W. Provine, eds., The Evolutionary Synthesis:
Perspectives on the Unification of Biology (Cambridge, 1980), p. 16.

54. E. Cochrane, op. cit.

55. It is also significant to note that several of these figures were
originally trained in biologyWatson, Pavlov, and Freud, for example.

56. D. Klein, A History of Scientific Psychology (New York, 1970).

57. W. Wundt, Outlines of Psychology (Leipzig, 1897), p. 319.

58. On this aspect of Freud's thinking, see Frank Sulloway, op. cit.,
pp. 92-94, 274-275.

59. Ibid., pp. 128-129.

60. S. Freud, Moses and Monotheism (New York, 1939), p. 128.

61. Jerome Bruner, writing in the preface to W. McDougall's Body and
Mind (Boston, 1961), p. xi.

62. W. McDougall, The Frontiers of Psychology (Cambridge, 1934), p.
200.

63. W. McDougall & J. Rhine, "Third Report on a Lamarckian
Experiment," British Journal of Psychology 24 (1933), pp. 213-235.  On
subsequent attempts to duplicate McDougall's results, see W. Agar et
al, "Fourth Report on a test of McDougall's Lamarckian experiments on
the training of rats," J. Exp. Biol. 31 (1954), pp. 307-321.

64. P. Fothergill, Historical Aspects of Organic Evolution (New York,
1953), pp. 261-262.

65. W. McDougall, op. cit., p. 177.

66. Quoted from a letter to Richard Bentley, dated February 25th,
1692/3.  In a previous letter to Bentley, Newton had objected as
follows: "You sometimes speak of gravity as essential and inherent to
matter.  Pray do not ascribe that notion to me, for the cause of
gravity is what I do not pretend to know."

67. J. Shapiro, "Genomes as smart systems," Genetica 84 (1991), p. 4.
The full statement is as follows: "Genomes function as true
intelligent systems which can be readjusted when conditions require.
We still lack testable theories to explain how these readjustments are
made."

68. J. Shapiro, "Genomes as smart systems," Genetica 84 (1991), pp.
3-4.

69. L. Margulis, Symbiosis in Cell Evolution (San Francisco, 1981).

70. John Maynard Smith, in a recent review of evolutionary theory,
offered the following observation: "A central idea in contemporary
biology is that of information.  Developmental biology can be seen as
the study of how information in the genome is translated into adult
structure, and evolutionary biology of how the information came to be
there in the first place."  E. Szathmary & J. Smith, "The major
evolutionary transitions," Nature 374 (March 16, 1995), p. 231.

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