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On Books
The Tangled Tale of Genes and Environment:
Moore’s The Dependent Gene:
The Fallacy of ‘‘Nature vs. Nurture’’
Susan M. Schneider
Florida International University
Nature–nurture views that smack of genetic determinism remain prevalent. Yet, the increasing
knowledge base shows ever more clearly that environmental factors and genes form a fully
interactional system at all levels. Moore’s book covers the major topics of discovery and dispute,
including behavior genetics and the twin studies, developmental psychobiology, and
developmental systems theory. Knowledge of this larger life-sciences context for behavior
principles will become increasingly important as the full complexity of gene–environment
relations is revealed. Behavior analysis both contributes to and gains from the larger battle for
the recognition of how nature and nurture really work.
Key words: nature–nurture relations, heritability, genetics, evolution, developmental systems
theory
91
92 SUSAN M. SCHNEIDER
mination of the number of fingers and seem. Consider the small number of
toes, say. But alternatively, given genetic diseases that are classified as
a ‘‘normal’’ genome, so-called environ- monogenic.1 Deriving from an ab-
mental determination of the number of normality in a single gene, these
fingers and toes could be claimed. constitute a very small fraction of
Consider the case of the teratogen diseases (Jablonka & Lamb, 2005). In
thalidomide, which frequently altered those monogenic diseases considered
this number during a tragic period in to be autosomal recessive (i.e., Men-
the 20th century. Further, smoking delian and not on the X or Y
during pregnancy is one of several chromosome), recessive homozygosi-
documented environmental risk fac- ty (two copies of the recessive form)
tors, although the evidence is correla- does not necessarily result in the
tional (see Man & Chang, 2006, who problematic phenotype. Nope. Put
demonstrated an epidemiological dose– technically, although those at genetic
effect relation). Indeed, at an elemen- risk can be highly likely to develop
tary level, a host of the right environ- the disease, the penetrance is never
mental factors must be present at the 100%, and it is sometimes consider-
right times and in the right places. Both ably lower (see Morange, 2001, for
genes and environmental factors are examples and commentary). Phenyl-
always necessarily involved. ketonuria (PKU), a classic genetic
Obviously, this conclusion in no disease of this type that is discussed
way diminishes the importance of the in The Dependent Gene, is character-
study of genetic contributions. Re- istic in that the severity varies despite
cent advances in genetics have been the same homozygosity—even con-
critical in demonstrating the often trolling for exposure to the problem-
Byzantine ways in which multiple atic amino acid that cannot be
genes and multiple environmental metabolized. ‘‘The evolution towards
factors interact. But headlines pro- seeing single-gene traits as versions of
claiming discoveries of the ‘‘gene for’’ complex traits has been under way
a wide range of human characteris- for some time,’’ noted Scriver and
tics, including personality and other Waters (1999, p. 267). They summa-
behavioral traits, require clarification rized numerous reasons for the var-
in a number of respects. iability in PKU outcomes, finding
First, a fundamental principle was that the major gene was just one of
well characterized in the early history many factors; indeed, ‘‘the whole
of genetics. As noted in The De- organismal phenotype is more than
pendent Gene, Sturtevant pointed out
the sum of the parts; it is an emergent
at the beginning of the 20th century
property’’ (p. 272). The Centers for
that, although a single gene had been
Disease Control’s panel of experts
found to be responsible for a differ-
ence in fruit fly eye color, other concluded in a general statement
factors being held as equal as possi- that, ‘‘As we acquire more knowledge
ble, that gene in no sense could be about the molecular basis of genetic
taken to code for eye color. Instead, disease, it becomes increasingly clear
eye color was the result of many that variable expressivity (i.e., modi-
genes and many environmental fac- fication of a genetic trait by other
tors. Moore suggests as an analogy genes or the environment) is the rule
the necessity of wheels and a chain in
order for bicycle pedals to operate for 1
Note the categorization difficulties with
forward motion. respect to the fuzzy set of nonmonogenic
Second, even with this important genetic diseases; note also other complica-
tions, such as the fact that the problematic
proviso, the simple single-gene single- allele for the monogenic disease sickle cell
trait systems popular in the media are anemia is actually beneficial in heterozygous
rare—and more complex than they individuals.
ON BOOKS 93
rather than the exception’’ (Burke et from simple. Most DNA, including
al., 1998, quoted in Moore, p. 230). human DNA, is what is known as
Third, in the version known as ‘‘junk’’—remnants from the past,
a phenocopy, PKU, like other genetic a proportion of which has turned
diseases, can develop in the absence of out to be regulatory. Cistrons, which
the known gene form (Gray, 2001; see constitute a tiny proportion of human
R. Moore et al., 2001, on Hunting- DNA, are those portions of a chro-
ton’s disease). Sometimes the prob- mosome that can actually code for
lematic mechanisms are identical and a protein. However, the cistrons are
sometimes they are different, but they not simple uninterrupted sequences of
result in either identical or nearly the relevant nucleotides; instead, they
identical symptoms. It will not come contain exons that actually hold the
as a surprise that the same symptoms sequence, intermingled with nucleo-
can be associated with either or both tides that generally appear not to code
genetic and environmental abnormal- for anything. Sometimes the exon is
ities in various combinations. In a small portion of the cistron. The
living systems, there are multiple cellular environment is critical for the
pathways to the same end. selection of the proper nucleotides to
Finally, a single gene commonly read. After all, all cells that constitute
influences many traits (pleiotropy). an organism contain the same ge-
Morange (2001) concluded, for ex- nome in their nuclei. Further, the
ample, that same cistron can be operated on in
different ways to code for different
There are no proteins specific to learning and proteins. A substantial proportion of
memory but rather proteins that, through their human DNA makes use of such
function as relays or transmitters, have been
harnessed by evolution in the development of alternative splicing. Jablonka and
cognitive processes. … What makes a process Lamb (2005) noted that one gene in
specific is not the nature of its molecular the chicken has been found to have
components (and thus the genes that code for 576 different splice versions—al-
these components) but the way they are used
and assembled in particular molecular path-
though, as is usually the case, these
ways and specific structures. (pp. 88–89) are minor variations of each other.
Finally, after the nucleotides are
These higher level operations neces- properly sequenced, the environment
sarily involve environmental factors. has long been recognized as essential
‘‘Genetically determined’’ could be for actual protein construction. For
seen as useful shorthand in some example, a protein’s shape, usually
cases, such as when a genetic feature critical for its function, depends on
like single-gene dominance or reces- environmental features as well as on
sive homozygosity often produces the sequence of specified amino acids.
given effects across a large range of This collection of findings means
‘‘normal’’ environments, just as, vice that the very definition of a gene can
versa, ‘‘environmentally determined’’ be less than straightforward. Al-
could for an environmental feature though the generic cistron usually
that often produces given effects qualifies, as Keller (2000) noted,
across a large range of ‘‘normal’’
genomes. But these simplifications The gene has lost a good deal of both its
can become problematic: They get specificity and its agency. Which protein
overgeneralized, and the fact that they should a gene make, and under what circum-
stances? And how does it choose? In fact, it
are simplifications can be forgotten. doesn’t. Responsibility for this decision lies
elsewhere, in the complex regulatory dynamics
Defining the Gene of the cell as a whole. It is from these
regulatory dynamics, and not from the gene
The very process by which genes itself, that the signal (or signals) determining
are said to code for proteins is far the specific pattern in which the final tran-
94 SUSAN M. SCHNEIDER
script is to be formed actually comes. (p. 63, changes are reversible, thus offering
quoted on p. 67 of Jablonka & Lamb, 2005) readier adaptability to changing con-
ditions than changes in the genes
In other words, environmental fac-
themselves.
tors are critical in determining what
These epigenetic mechanisms are in
protein-coding exons get read from turn responsive to more molar-level
a cistron, when, and how often. Thus, environmental factors. But so of
the very concept of a gene requires course is the genome itself. Chemicals
the environment. As Moore puts it, and electromagnetic emissions are
among the environmental factors well
Such contextual dependence renders untena- known to be capable of altering
ble the simplistic belief that there are coherent,
long-lived entities called ‘‘genes’’ that dictate DNA directly. Environmental and
instructions to cellular machinery that merely behavioral factors routinely modify
constructs the body accordingly. The common gene expression and activity (see
belief that genes contain context-independent Gottlieb, 1997, 1998, for numerous
‘‘information’’—and so are analogous to documented ways). Immediate early
‘‘blueprints’’ or ‘‘recipes’’—is simply false.
(p. 81) genes, for example, are activated by
environmental signals. Genetic muta-
Strictly speaking, then, as Gray tions even appear to be induced when
(1992) concluded, ‘‘a gene can only and where needed to some extent (see
be functionally defined in a specific Jablonka & Lamb, 2005, for an
developmental context’’ (cited in extensive discussion). Stress is one
Moore, p. 81). of the factors documented to result in
mutations in the DNA (and some
Genes, Epigenetics, and controversy exists over whether that
Epigenetic Inheritance phenomenon has been selected for or
is simply a side effect). In any event,
The cellular-level mechanisms in- as Moore notes, ‘‘our daily experi-
volved in these operations are epige- ence of stress directly impacts the
netic, meaning that they entail non- activity of our genes’’ (p. 139). Com-
genetic factors that are inherited pensations for these stress effects can
themselves or that affect genetic in- occur by an adjustment of the ex-
heritance and gene expression. For pression of the same gene or in other
example, DNA methylation, which ways (e.g., Francis, Diorio, Plotsky,
does not affect the genotype, reduces & Meaney, 2002): Again, multiple
the likelihood of gene expression. pathways exist toward the same end.2
Methylation patterns can themselves Cellular epigenetic mechanisms are
be inherited. If the capacity to pro- among the important mediators.
duce proteins for a needed function is Several types of epigenetic inheri-
present in the genome, it can be tance systems have been discovered,
unmasked through epigenetic means. including RNA interference (subject
(A dramatic demonstration was Kol- of a recent Nobel prize), prions, and
lar & Fisher’s, 1980, induction of DNA methylation and other forms of
teeth from bird tissue in vitro.) The chromatin marking (i.e., marks on
resuscitated gene can then be avail-
able once again for natural selection 2
In the same way, gene knockouts can have
to act on. Epigenetic mechanisms surprising effects, sometimes even causing
improvements in functioning although the
also have large effects on the DNA protein coded for had been thought to be
that helps regulate the protein-coding critical (e.g., Morange, 2001). In this regard,
genes, such as the transposons (so- Jablonka and Lamb (2005) refer to ‘‘the
called jumping genes that constitute dynamic regulatory structure of the network’’
(p. 63) as a potent force for stability. Moore
a large proportion of the mammalian has been criticized for failing to discuss the
genome). And, as Jablonka and knockout gene studies, but they can actually
Lamb (2005) noted, epigenetic be taken to bolster his case.
ON BOOKS 95
the materials that form the chromo- ures like 70% have been produced for
somes). As contrasted to the genetic the heritability of IQ; the heritability
system, the epigenetic inheritance of autism has also received a high
system transmits phenotypes, not estimate. What do these numbers
genotypes, a feature it shares with really mean?
behavioral inheritance systems (see
below). Further, changes during an Understanding Heritability
organism’s own lifetime are some- To begin, Moore summarizes a
times inherited, in Lamarckian fash- famous illustration by Lewontin
ion, across generations of organisms (1970). If seeds varying in genetic
as well as of cells within an organism constitution are raised in identical
(Jablonka & Lamb, 1995, 2005). environments, any differences among
Astonishingly, one researcher turned the plants, such as height, must be
some of the cilia of a paramecium due to genetic variation. Thus, heri-
inside out—and the change was tability for height (or any other trait)
inherited (Beisson & Sonneborn, must be 100%. If the same seeds are
1965, also cited in Jablonka & Lamb, sown in identical hostile environ-
2005, p. 122). Such environmentally ments, all the plants are much
induced changes can sometimes be- shorter, but height differences among
come assimilated in the genome, a fact them must still be due to genetic
known since Waddington’s classic variation, and heritability remains
fruit fly experiments in the 1940s 100%. Yet, the differences between
(see Avital & Jablonka, 2000; also see these two groups obviously depend
Moore, p. 202). These mechanisms on the environments. And, whatever
do not act only in single-celled the heritability, plants need soil,
organisms. Pavelka and Koudelova water, and sunlight to grow.
(2001) found that Mediterranean
Moore continues the analogy with
flour moth larvae with a mutation
the example of cloned seeds (seeds
for short antennae developed nor-
with identical genes) raised in envir-
mal-length antennae as adults, if
onments that are not identical. In this
raised at a higher incubation temper-
case, any height or other trait differ-
ature than normal during a sensitive ences must be due to environmental
period. Their offspring for several differences, so heritability is 0. But
generations retained this feature, de- genes are obviously necessary. For
spite the short-antennae genotype, the same trait in the same species,
and despite being raised at the normal then, heritability can vary through-
incubation temperature, with epige- out its range as a function of
netic inheritance mechanisms consid- circumstances. (Indeed, the heritabil-
ered the most likely cause. Examples ity of IQ has long been known to be
in vertebrates also exist (e.g., mouse substantially lower in children than in
health characteristics and coat color; adults, e.g., Block, 1995.)
see Jablonka & Lamb, 2005).
Two examples drawn by Moore
from Block (1995) bring home the
HERITABILITY
point. First, the number of human
Inheritance is complex, and Moore’s fingers and toes has very low herita-
deconstruction of heritability shows bility. Variability in digit number is
how simplistic and misleading the largely accounted for by accidents or
usage of that construct has often disease—environmental factors, not
been. Heritability is defined as the genetic variation. As discussed pre-
proportion of trait variation associ- viously, the teratogen thalidomide
ated with corresponding genetic var- provides one example: When it acts
iation—in a particular population prenatally, despite a ‘‘normal’’ ge-
under particular circumstances. Fig- nome, an abnormal number of digits
96 SUSAN M. SCHNEIDER
can result. Second, the wearing of tical twins raised in different environ-
earrings in 1950s America had high ments may share a trait outcome not
heritability: Only females used to be because of their shared genes, but
likely to wear earrings then, explain- because of similar or different fea-
ing the genetic correlation. But cul- tures of their different environments,
tural factors were clearly as critical features that might have produced
then as they are now for this behav- the same outcome regardless of a wide
ior, now that its heritability must be variety of genetic differences. Along
lower. So, although heritability these lines, being raised in the same
sounds like it quantifies the degree family does not mean that environ-
to which a trait itself is determined by ments do not vary in many significant
genes, it does not. (And of course it ways. Just one such difference can be
could not: Genes and environmental enough to create a large and long-
factors are both always necessary; lasting effect on a trait, a point John
recall Moore’s example of the bicycle B. Watson made many years ago. As
pedals for forward propulsion only behavior analysts know, individual-
in conjunction with other essential ized operant and classical condition-
parts.) ing histories are critical in the de-
velopment of behavioral patterns and
Limitations, Confusions, and characteristics.
Confounding Variables For the purposes of heritability
estimates, genes and environments
This is just the beginning of the can be directly controlled only for
confusions concerning this correla- plants and some nonhuman animals,
tional construct. Heritability esti- and even then, these efforts often fail.
mates statistically apportion sources In a Science article (Crabbe, Wahl-
of variation in traits, but they apply sten, & Dudek, 1999), for example,
only to the specific populations and mice from the same genetic strains
contexts from which they are derived. were raised in different laboratories
They cannot be generalized to other under environments rigorously con-
populations or circumstances without trolled to be as similar as possible.
extra empirical evidence. And if the On a number of behavioral tests,
original context varies—if environ- however, different laboratories found
ments are sometimes similar and different results for the same genetic
sometimes different in ways that strain, differences sometimes bigger
affect the trait—the estimates them- across laboratories than across
selves are confounded. Heritability strains. (The short-term, less-than-
estimates apply only to groups, and precise nature of the tests, such as
are inherently inapplicable to indi- open field and maze, as contrasted to
viduals in any sense. And they do not longer term behavior-analytic oper-
imply causation. As Moore notes, all ant and respondent assays, makes
of these important limitations have these results less surprising;3 also see
been frequently ignored or mini- Francis, Szegda, Campbell, Martin,
mized. & Insel, 2003.)
Consider also a pair of identical For humans, bombarded by rich
twins reared in different environ- and varied experiences every day,
ments. If analogous plant clones many of the environmental factors
grow to the same height in different
environments, this identical outcome 3
Encouragingly, Crabbe et al. conclude by
cannot be concluded to be ‘‘pro- noting that ‘‘increased communication and
grammed by the genes’’ in any sense: collaboration between the molecular biologists
creating mutations and behavioral scientists
Lack of sun in one location may be interested in the psychological aspects of
matched in effect by poor soil in behavioral testing will benefit both groups’’
another, for example. Similarly, iden- (p. 1672).
ON BOOKS 97
extent to which genes control the trait’s a reinforcer later, and can perhaps
development. (Moore, p. 185) contribute to alcoholism (e.g., Spear
Farewell (again) to the percentage & Molina, 2005; this particular re-
game. search line is not cited in The De-
pendent Gene). Here again, confusion
As Moore points out, the detection
can arise over genetic and nongenetic
of such nonobvious contributors
familial inheritance patterns.
requires special care. Mother rats’
licking of male preweanlings has been
Behavioral Inheritance
shown to be essential for the later
development of normal sexual behav- Developmental work has comple-
ior (C. Moore, 2003). However, mented behavioral work in docu-
separating the pups from their moth- menting nongenetic inheritance mech-
er after weaning, raising them in anisms in addition to the more mole-
social isolation, and observing nor- cular epigenetic ones discussed pre-
mal sexual behavior might be taken viously. For example, it has long
to suggest that the environment is been known in humans and other
unimportant, which is clearly far mammals that acquired immunity
from the case. Many such examples can be transmitted nongenetically,
of nonobvious environmental contrib- through breast milk and the placen-
utors are now known to exist (see, ta. Similarly, when a female Mon-
e.g., Lickliter & Honeycutt, 2003). golian gerbil embryo is positioned
Experience is critical for develop- near brothers rather than sisters, she
ment in myriad ways, and Moore is exposed to more testosterone, and,
notes research showing that corre- like her male siblings, is likely to be
sponding brain plasticity is now licked more than female-positioned
known to be higher throughout the females (Clark, Bone, & Galef, 1989;
lifespan than had been thought. The Clark, Karpluk, & Galef, 1993).
fantastic chimeras created by embry- Later behavioral effects include
ologists who combine parts of differ- greater aggression and the ability to
ent creatures have demonstrated how hold larger territories. Such female
the environment, not the genes, gerbils tend in turn to have male-
determines which cells become parts dominated litters, so their daughters
of what organs, and just how plastic show the same patterns, thus pro-
that process is. ‘‘We are standing and viding another illustration of non-
walking with parts of our body which genetic inheritance. Behavioral mech-
we could have used for thinking if anisms are involved, and the extra
they had been developed in another licking provides an excellent exam-
position in the embryo’’ (Spemann, ple. Further, as discussed previously,
quoted in Moore, p. 87). similar extra licking of male pups by
Moore, an infancy researcher him- rat mothers was demonstrated to be
self, focuses especially on perinatal critical for later male sexual behav-
development, the source of an explo- ior. This behavior has been shown to
sion of news over the past few be caused by testosterone or associ-
decades. One phenomenon is fetal ated hormones in the male rat pups’
programming, a lifelong predisposi- urine, which act as a reinforcer for
tion to obesity caused by poor the mothers’ licking (C. Moore, 1982,
maternal nutrition at a particular 1995).
prenatal stage. Of special interest to Cross-fostering studies, in which
behavior analysts, Spear and his young of one genetic strain are reared
colleagues have shown that placental by mothers of a different strain, are
or mammary exposure to ethanol (at especially useful in studies of gene–
levels far below those for fetal environment inheritance relations.
alcohol syndrome) establishes it as Cierpial and McCarty (1987) used this
ON BOOKS 101
approach to show that rats of the passed along rather like genes (and
spontaneously hypertensive (SHR) the essential cytoplasm containing
genetic strain do not show the SHR the genes):
behavior pattern if raised by non-
SHR mothers (see also Ressler, 1966, To the extent that we cannot help but develop
discussed by Moore; C. Moore, in environments that are similar in important
Wong, Daum, & Leclair, 1997; Suomi, ways to the environments in which our parents
developed, the legacy we receive from our
1999). Integral once again were be- parents includes both our genes and aspects of
havioral mechanisms similar in some our developmental environments. (Moore,
ways to the differential maternal p. 174)
handling discovered by C. Moore
(Cierpial, Murphy, & McCarty, 1990). Evolutionarily speaking, both genes
Operant behavior comes even and critical features of environments
more to the forefront in the social are and must be reasonably stable
learning that is an obvious behav- across generations. Second, as Moore
ioral inheritance mechanism. Berman points out, natural selection does not
(1990) noted likely operant involve- act directly on genes, but on pheno-
ment in the maternal parenting styles types. Phenotypes are produced and
that tend to be passed down from modified by both genes and environ-
mother to daughter for generations in ments, and behavior principles have
rhesus monkeys (see Fairbanks, 1996, an important role. Evolution might
and Suomi, 1999, for related re- even be considered to proceed by
search). For example, access to an lasting phenotypic changes regardless
infant sibling is reinforcing for most of whether there is an accompanying
females, and maternal rejections of change in the genome, a controversial
the infant can be discriminative proposal made by Gottlieb (Moore,
stimuli signaling an opportunity for p. 201). These lines of thought are at
access. Attention to the mother’s the heart of the integrative, empiri-
parenting of the sibling is sometimes cally based approach to nature–nur-
reinforced by access to the mother as ture relations known as developmen-
well. Berman suggests that such tal systems theory.
stimulus control facilitates learning
of a parenting style through imitation Developmental Systems Theory
(which of course involves operants; The Dependent Gene is one of the
see, e.g., Chase & Masia, 1997). first trade books on developmental
Observational learning is also critical systems theory, which encompasses
for the transmission of foraging all the research areas bearing on
techniques. An impressive variety of nature–nurture relations. Behavior
such behavioral inheritance mecha- analysis is eminently consistent with
nisms across the animal kingdom is this approach, one that makes the
documented in Animal Traditions: role of environmental factors like
Behavioural Inheritance in Evolution behavior principles explicit. The very
(Avital & Jablonka, 2000), a work in title of a recent edited work in this
which operant involvement is explic- tradition is significant: Cycles of
itly recognized. Contingency (Oyama, Griffiths, &
The evolutionary implications are Gray, 2001). Moore’s book provides
significant. Moore only footnotes the an excellent introduction. A more
Baldwin effect—the idea tracing back technical work of epic scope is
to Lamarck and Darwin that behav- Jablonka and Lamb’s (2005) Evolu-
ior can initiate evolutionary change tion in Four Dimensions: Genetic,
(see Avital & Jablonka, 2000; C. Epigenetic, Behavioral, and Symbolic
Moore, 2003; Schneider, 2003). How- Variation in the History of Life. Other
ever, he emphasizes two key associ- notable recent books that can rea-
ated insights. First, environments are sonably be grouped under the de-
102 SUSAN M. SCHNEIDER
twin study. Journal of the American Acad- research, and clinical applications (pp. 181–
emy of Child and Adolescent Psychiatry, 45, 197). New York: Guilford.
691–699. Suomi, S. J. (2002). How gene-environment
Schneider, S. M. (2003). Evolution, behavior interactions can shape the development of
principles, and developmental systems: A socioemotional regulation in rhesus mon-
review of Gottlieb’s Synthesizing Nature- keys. In B. S. Zuckerman, A. F. Zucker-
Nurture: Prenatal Roots of Instinctive Be- man, & N. A. Fox (Eds.), Emotional
havior. Journal of the Experimental Analysis regulation and developmental health: Infancy
of Behavior, 79, 137–152. and early childhood (pp. 5–26). New Bruns-
Scriver, C. R., & Waters, P. J. (1999). Mono- wick, NJ: Johnson & Johnson Pediatric
genic traits are not simple: Lessons from Institute.
phenylketonuria. Trends in Genetics, 15, Suomi, S. J. (2003). Gene-environment inter-
267–272. actions and the neurobiology of social
Spear, N. E., & Molina, J. C. (2005). Fetal or conflict. Annals of the New York Academy
infantile exposure to ethanol promotes of Sciences, 1008, 132–139.
ethanol ingestion in adolescence and adult- Todd, J. T., & Morris, E. K. (1992). Case
hood: A theoretical review. Alcoholism: studies in the great power of steady mis-
Clinical and Experimental Research, 29, representation. American Psychologist, 47,
909–929. 1441–1453.
Suomi, S. J. (1999). Attachment in rhesus West-Eberhard, M. J. (2003). Developmental
monkeys. In J. Cassidy & P. R. Shaver plasticity and evolution. Oxford: Oxford
(Eds.), Handbook of attachment: Theory, University Press.