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Lamarckian Effect: Evidence of the Genetic Inheritance of Adopted Features in the Neuropsychological Literature.

[abstract only] "Institute of Zoology, University of Genoa, Italy. The cell property of processing the environmental information is called sensitivity and referred to a "sensitive" cell system in which physical signals and specialized molecules work in relation to peculiar cell structures. It is suggested that, in addition to its role in cell behaviour, the sensitive system performs a basic role in morphogenesis in all cells. It interacts with the genetic system in gene expression and regulation directly or indirectly through epigenetic processes. Epigenetic inheritance, adaptive mutations and genetic assimilation allow environmental information to be embodied in new traits, thereby becoming relevant in evolution. This is consistent with an evolutionary mechanism with two alternate and recurrent phases: the first, epigenetic and Lamarckian at cell and organism levels and the second, selective and Darwinian, at population level." (Sara M (1996) A "sensitive" cell system. Its role in a new evolutionary paradigm. Riv Biol 89(1):139-56)

"To put the matter as succinctly as possible, no case has yet been satisfactorily proved in which, as a result of external factors, the development of an animal has undergone a modification and in which these external factors have become internal and transmitted, so that the same modification has come to be invariably produced in all the subsequent ontogenies of descendant animals without the necessity for the external factors which orginally evoked the modification. Until such a case has been proved, it cannot be believed that the effects of external factors and of use and disuse on body or mind are transmitted or play any part in ontogeny in subsequent generations. Somatic induction, or the transmission of the effects of use and disuse, constitute the kernel of the Lamarckian point of view; and it is curious that while we still lack evidence, viz. direct induction, though of course he was prophetic and correct in rejecting the view that direct induction can produce an adaptive inherited response to the environmental stimulus. It would be very convenient if it were possible to accept an explanation of the origin of internal factors and of their adaptive nature on the lines of Lamarck's hypothesis of the effects of use and disuse, but in the present state of knowledge it is not possible. It is necessary to adopt a humbler position and admit that the causes of origin and change in the genetical factors of organisms are unknown. Once they have arisen or changed (mutated), selection plays an all-important part in moulding their effects." (de Beer, G. R. (1951) Embryos and Ancestors. Clarendon Press: Oxford. p. 15-6)

"The Lamarckian hope that a discovery is just around the corner, that somewhere, perhaps in some recondite corner of the immune system, acquired characteristics will turn out to be inherited, is in vain. In our world or epigenetic recipe embryology, the inheritance of acquired characteristics is impossible." (Cronin, Helena (1992) The Ant and the Peacock: Cambridge Univ. Press, Cambridge p. 44)

"High growth rates have independently been suggested as one of several adaptations that differentiate folivores from nonfolivores (Leigh, 1994a), accompanying dental and gut specializations that facilitate consumption of high-fiber resources by folivores (Kay, 1984; Fleagle, 1988). Elevated growth rates may confer additional advantages to gorillas and other folivores, including metabolic advantages to early attainment of large size and early maturation of alimentary system, especially if the gut scales with positive allometry during ontogeny. (see Leigh, 1994a). Therefore, we suggest that high growth rates in gorillas are partly related to folivory through physiological advantages of rapid growth and reduction in ecological risks. Obviously, we cannot rule out the possibility that a derived size increase is related to increase folivory in gorillas. Ecological risk in gorillas is further reduced relative to common chimpanzees because they utilize large and ubiquitously distributed food patches (Wrangham, 1986). Enhanced growth rates might also confer reproductive advantages to females because early maturation (or early attainment of an "optimal" female adult size) is not comstrained by slow growth. (Rubenstein, 1993; Pagel and Harvey, 1993). We must note that lowland gorillas (studied here) are likely less folivorous than mountain gorillas (Tutin and Fernandez, 1985,1993), which serve as the basis for most of our information aobut Gorilla adapatations (Watts, 1984, 1985, 1990, 1994). However, levels of fovivory in lowland gorillas are probably sufficient to reflect adaptations that characterize other folivorous primates (over 40-45% of annual time spent feeding on folivorous resources seems to be sufficient to reflect this adaptation {Leigh, 1994a}. Thus, the resent analysis suggests that Janson and van Schaik's (1993) model had predictive power for explaining why gorillas grow more rapidly than other African apes, and contributes to an understanding of adult size differences between genera." (Leigh, S.R. & Shea, B.T. (1996) Ontogeny of body size variation in African apes. American Journal of Physical Anthropology 99(1): pp. 60-61)

"Another example is much more common, but less acknowledged. It is the case of cell differentiation in embryonic development. Once a cell differentiates by responding to an environmental factor (i.e. an inductor substance), it usually passes its characteristics to descendants. (Alberts et. al. 1989, Blau, 1989). Again, after the acquisition of the differentiated character, there is hereditary transmission of it." (Aboitiz, F. (1992) Mechanisms of adaptive evolution. Darwinism and Lamarkism restated. Medical Hypothesis 38(3): pp. 196)

"In the light of modern genetics, it is presently considered that IAC is practically an impossibility. In metazoans, the germline and the genetic material are supposed to be isolated form perturbations coming from the environment. Thus, ontogentic changes may not effect heredity. This assumption seems correct for metazoans, in whom the majority of phenotypic traits is indirectly inherited. However, as already mentioned, interitance of acquired characteristics becomes more plausible in cases in which direct inheritance makes a significant portion of the phenotype, as in cell division. If, for example, a cytoskeletal feature is modified during a cell's ontogeny, this is likely to be inherited by descendants. This point was referred to as the transgenerational dimension of modifications to directly inherited characters." (Aboitiz, F. (1992) Mechanisms of adaptive evolution. Darwinism and Lamarkism restated. Medical Hypothesis 38(3): pp. 196)

"As in Bolitoglossa occidentalis, it may transpire that a single heterochronous disturbance delaying growth and development and making the whole epigenetic system more vulnerable to stochastic factors and environmental perturbations will be a suffecient explanation of the pathogenesis of Down syndrome." (Opitz, John M. & Gilbert-Barness, Enid F. (1990) Reflections on the pathogenesis of Down syndrome. American Journal of Medical Genetics 7: pp. 49)

"They showed in Table I and II and Figs. 1 and 2. Alcohol patients showed significantly lower testosterone levels than controls. The circadian rhythm was lost after withdrawal but it recurred on third day and progressively increased until full recovery, i.e. the difference between hormone levels at 9 and 21 h increased slowly, reaching nearly the level of signification on day 15. FSH and LH levels were signifantly higher in alcoholics at day 1, and the levels did not vary significantly during the 15 days of the study. A strong relationship existed between levels of FSH and LH (r = 0.56, P< 0.0001), but not between these hormones and testosterone. Prolactin levels significantly increased during the 15 days of observation, being in the interior limit of the normal range at the beginning." (Castilla-Garcia, A., Santolaria-Fernandez, F.J., Gonzalez-Reimers, C.E., Bastita-Lopez, N., Gonzalez-Garcia, C., Jorge-Hernandez, J.A., & Hernandez-Nieto, L. (1987) Alcohol-induced hypogonadism: reversal after ethanol withdrawal. Drug and Alcohol Dependence 20(3): pp. 257)

"In our study, testosterone levels progressively returned to normal along the 15 days of the observation period. In addition, the circadian rhythm, lost at the beginning of the study --- a feature also reported by Gordon (2) and Bertello(22) --- became evident again in the last days of the observation period. These data suggest that hypogonadism seems to be functional in nature. .... Prolactin has been found to be elevated in alcholics after a week of abstinence (13,23), and decreased during the hangover phase, with blockade of the prolactin response to TRH (24). In this connection it has been argued that ethanol withdrawel may lead to an excessive dopaminergic activity, and, as a consequence, to a low release of prolactin (24). In our study, serum prolactin levels-especially those obtained at 21 h --- recovered progressively from the low limit of the normal range observed in the first days to the higher levels at the end of the observational period." (Castilla-Garcia, A., Santolaria-Fernandez, F.J., Gonzalez-Reimers, C.E., Bastita-Lopez, N., Gonzalez-Garcia, C., Jorge-Hernandez, J.A., & Hernandez-Nieto, L. (1987) Alcohol-induced hypogonadism: reversal after ethanol withdrawal. Drug and Alcohol Dependence 20(3): pp. 259)

Chart (book wall) comparing testosterone levels of Chinese, Japanese, African and Whites, all USA. Whites had lowest level of the four, contradicting shift theory expectations. Tests need to be done of ethnic groups within their cultures, that stress increases testosterone levels could heavily influence the figures in this study; the measurements may be noting the relative stress of community ties and socioeconomic status. (Wu, A.H., Whittemore, A.S., Kolonel, L.N., John, E.M., Gallagher, R.P., West, D.W., Hankin, J., Teh, C.Z., Dreon, D.M., & Paffenbarger, R.S.Jr. (1995) Serem androgens and sex hormone-binding globulins in relation to lifestyle factors in older African-American, white, and Asian men in the United States and Canada. Cancer Epidemiology, Biomarkers & Prevention 4(7): pp. 738)

"After adjustment for age, Quetelet's index, and physical activity, levels of testosterone (total, free, and bioavailable) were higher for Asians born in Asia versus for those Asians born in North America, whereas levels of SHBG and DHT were similar in both groups (Table 4). On the other hand, the DHT: testosterone ratio was lower in Asians born in Asia compared to those born in North America. However, none of the differences between Asians born in North America and those born in Asia were statistically significant." (Wu, A.H., Whittemore, A.S., Kolonel, L.N., John, E.M., Gallagher, R.P., West, D.W., Hankin, J., Teh, C.Z., Dreon, D.M., & Paffenbarger, R.S.Jr. (1995) Serem androgens and sex hormone-binding globulins in relation to lifestyle factors in older African-American, white, and Asian men in the United States and Canada. Cancer Epidemiology, Biomarkers & Prevention 4(7): pp. 739)

"Stenuous physical activity may result in lower levels of testosterone (26-28). Smoking may inhibit production of estrogens in women (29) but data on its effect in men are not consistent (30,31). In a study of Caucasian upper middle class men in a California retirement community, smokers showed significantly higher serum estrogen levels than did nonsmokers (30). However, in the Multiple Risk Factor Intervention Trial, serum estrogen levels in men were not associated with total amount of cigarettes smoked (31). Levels of testosterone were higher among men who smoked compared withmen who did not in both studies (30,31). Heavy alcohol use may increase estrone production and, thus, increase conversion of androgens to estrogens (32, 33). The present findings suggest higher total testosterone levels in smokers compared to nonsmokers, but there were few consistent associations between physical activity, alcohol use, and androgen levels. In this study, men with a self-reported history of a benign prostate condition showed a signigicantly higher DHT: testosterone ratio, a marker of 5a-reductase activity (34), compared to men without such history. This observation is compatible withthe hypothesis that the development of benign prostate (35). .... The present study of older men, most of whom were 60 years or older, found lower testosterone levels in whites compared to African-Americans, similar to studies conducted in young college men and Vietnam veterans aged 31-50 years. However, there is little evidence from this and other studies that testosterone levels in Asians residing in Asia or in North America are lower than those in whites. On the other hand, the strongest support for ethnic differences in androgen levels in this study was bsed on the DHT:testosterone ratio, which was highest in African-Americans, intermdiate in whites, and lowest in Asian-Americans, reflecting the prostate cancer incidence rates in these groups." (Wu, A.H., Whittemore, A.S., Kolonel, L.N., John, E.M., Gallagher, R.P., West, D.W., Hankin, J., Teh, C.Z., Dreon, D.M., & Paffenbarger, R.S.Jr. (1995) Serem androgens and sex hormone-binding globulins in relation to lifestyle factors in older African-American, white, and Asian men in the United States and Canada. Cancer Epidemiology, Biomarkers & Prevention 4(7): pp. 739-40)

"First, both physical and psychological stress have been consistently found to depress T (Mason, 1968; Kreuz et al., 1972; Opstad & Aakvaag, 1982). Second, aggressive or competitive/dominance encounters generally raise T levels, which also reflect the outcome of the encounter: T levels are elevated in winners and relatively repressed in losers (Rose et al., 1975; Elias, 1981). ... Generally, sustained moderate-to-intense exercise appears to elevate T, although data from studies have been equivocal (Kuoppanalmi et al., 1980)." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 450)

"In Western clinical populations, a marked diurnal variation in serum T levels has been reported, with evening values averaging around 30% lower than morning values (Nieschlag, 1974). Baseline values of !Kung hunters show similar diurnal variation." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 452)

"Evolutionary arguments for the importance of hunting in shaping foundations of men's behavior have tended to focus on its aggressive, competitive and stressful components. Its basic nature as a form of effort, or work, has been less frequently considered. The data suggest that !Kung men do not experience subsistence hunting as mere stress, which depresses T levels, for its main effect was an increase in evening values. On the other hand, no interaction of hunt outcome and T levels were observed, in contrast to reports that T reflects success and failure in certain aggressive and dominance interactions. Rather, these data are most congruent with reported effects of moderate prolonged excercise." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 456)

"We conclude that subsistence hunting was associated with significant changes in testosterone patterns in !Kung hunters, due primarily to the maintenance of T concentrations during hunting days that is reflected in elevation of evening values. Diurnal variation in circulating T was hence markedly attenuated. It is also notable that shifts in T level were a consequence, not a primary cause, of the behaviors observed here; hormone changes followed and supported, not preceded, hunting activity. Studies of human behavioral biology might consider the ways in which the dominant context of human evolution, the hunting-gathering niche, may have selected for efficient, appropriate responses to subsistence-related effort." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 456)

"Readers will question why I should attempt reviving such biological heresy long since thought buried. The seasons lie not in Darwin's concept of selective survival but with the neo-Darwinian reliance on two notions which have guided thinking on the subject for about a century. Currently these notions are (a) that the locus of all meaningful phylogenetic changes (in multicellular organisms) occurs only in the germline genes of the gonads, (b) that the genes in the gonads are resistant, with the exception, for example, of ionizing radiation, to most direct influences of the contemporary environment. I do not feel these premises, when coupled to the concept of selective survival, help explain the phenomenon of evolution as we perceive it." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 2)

"The other point, which Mayr's summary illuminates, is that Darwinian selection as perceived in the current paradigm is basicly a negative selection theory. Variant individuals arise in a population and those which by chance are unfit for survival are eliminated by the environment. This ordering process
provides a rational mechanism for guiding evolutionary change to some extent in large populations of individuals, but I feel it is a rather crude conceptual device and unlikely to be the final word on the evolution of multicellular organisms: to enrich evolutionary theory I think it will be necessary to add the complementary, yet opposing, concept of a positive selection force (feedback) also supplied by the environment." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 45)

"If the central dogma is correct, what do we do with Weismann's doctrine? Can it be replaced with something more useful and rational? I am aware of no direct observations which unequivocally prove that genetic information (DNA, RNA) cannot pass, in principle, from differentiated body cells to ova or sperm. There is also no good evidence that it can. I will stick my neck out and predict that future research on the endogenous C-type RNA viruses will very soon supply evidence that C-type RNA viruses can (a) "capture" genes in somatic cells (probably mRNA), (b) cross the tissue barriers partitioning the gametes in the gonads, (c) infect ova or sperm (or travel as extracellular particles in the seminal fluid, infecting the egg around the time of fertilization), and (d) by a process involving reverse trascriptase (RNA-dependent DNA-polymerase), synthesize a DNA copy allowing the integration of these captured somatic genes ("normal" or mutated) back into their appropriate positions in the germline DNA. This step is a crucial feature of the model because it demands a high degree of integration fidelity as to where "new" gentic information is to be positioned to satisfy the four dimensions of future development." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 48)

"The hypothesis states that new adaptive processes begin within the somatic cell populations of individual organisms. Environmental stimuli select and order genetic mutations occurring by chance in somatic cells of a given tissue or organ; if the selection stimulus is intense and chronic, there is an increased probability that these new "adaptive" genes are captured by the appropriate endogenous viral vector and integrated back into their homologous Medelian loci in the germline DNA of that individual. In this way, the organism both adapts to its own environment and anticipates, to some extent, the future environment of its offspring: allowing them to adapt instictively, if this selection process continues. The synthesis of this hypothesis is achieved by fusing two widely acknowledged theories, both of which enjoy considerable experimental support: (1) the clonal selection and somatic mutation theory of antibody production and (2) the proto-virus and provirus hypothesis of the endogenous RNA tumor viruses." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 60)

"It is very difficult to decide how far changed conditions, such as of climate, food, etc., have acted in a definite manner. There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence." Charles Darwin, The Origin of Species, 6th addition, 1872, p. 139. (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.viiii)

"I have endeavored to integrate the ever-growing knowledge from endocrinology in the discussions of all animal groups concerned. The result shows that structural changes that occur during the proximate process appear to result form the interaction of the endocrine mechanism inherited from the ancestors and new enviroment stimuli. Hence the structural modifications that resulted have, at least initially, nothing to do with natural selection. Often the latter is shown to have contributed merely to the improvement of the endocrine mechanism for such changes." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.x)

"Many environmental factors, such as food, temperature, photoperiod, and population density, affect the endocrine activity. Of these, nutrition is most fundamental, because the organs or cells that produce and release hormones must grow to be functional. These environmental factors induce production and/or release of hormones." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp. 7)

"Adaptive alterations of physiological functions of various organs and structures suited for terrestrial life have also accompanied the metamorphosis of externally visible structures. Frieden and Just (1970) summarized adaptive changes during metamorphosis as follows:
1) Tail regression and limb development leading to more powerful locomotion of land.
2)The shift form ammonotelism to urolelism reflecting the change in environemental water availability.
3) Change in hemoglobin reflecting the greater availability of ozygen in the frog.
4) Increase in serum proteins, particularly serum albumin, reflecting homeostasis and maintenance of the circulatory volume.
5) Changes in digestive enzymes and in intestinal design reflecting the necessary adjustment to a signigicant alternation of the diet."
(Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.11)

"Despite some apparent exceptions, it now has to be concluded that an environmentaly modified phoenotype could become heritable through the process of gentic assimulation, remaining at phase 1 or 2, or 3 or tha latter. As I (Matsuda 1982) have discussed, the widely held belief that environmentally modified phenotypes are not heritable (and hence without evolutionary future) is misleading and it must be dismissed. The reason for the prevalence of this belief appears to have been a series of proposals and discoveries of important biological concepts that have confused biologists. Most prominently, the so-called Weismann theory was in accord with the idea of impossibility of inheritance of externally induced phenotypic changes. The theory states that only the germ plasm is continued from generation to generation, and therefore somatic (somatogenic of Weismann) modifications acquired during development by external influence cnnot affect the germ plasm, and hence cannot be inherited. Against the possibility of transmission of somatic modification to the germ plasm, Weismann (1892, p. 393) asserted "we should have to assume the presence in all parts of the body a definite track along which somatic variation might be transferred back to the germ cells, in the germ plasm of which it would produce a corresponding change." This assertion was consistent with the Mendelian law of heredity that was discovered soon after, and the two theories together contributed to the development of neo-Darwinism, in which the external (environmental) influence on development is irrelevant to evolution. Further, Weismann's theory was also consistent with the central dogma of modern molecular genetics in which the relationship between the genotype and phenotype (i.e., the DNA to RNA to protein to phenotype sequence) is unidirectional and there is no mechanism by which the process could be reversed, and later repudiations of the "inheritance of acquired characters" (Dobzhansky 1970, Mahr 1976, Ayalo 1977) were based on this dogma. However, in these repudiaitions, as Matsuda (1982) pointed out, no distinction was made between the kinds of external agents causing modifications. In this connection, it is very important to point out here that Weismann (1892) acutually recognized two kinds of somatogenic variations, namely, injuries and functional variations, and the variations depending on the so-called "influence of environment" which included mainly climatic variations. As his discussion shows, his refutation of the "inheritance of acquired characters" applied to the first two categories only. As this work abundantly shows, environmentally acquired characters can become heritable through the process of gentic assimulation, without requiring the reversal of genetic transcription and translation. Weismann found, with regard to the climatic influence, that when the pupae of the German form of a lycaenid butterfly Polymmatus phlaeas was exposed to much higher temperatures, none of the emerged adults resembled the darkest form of sothern veriety eleus. Further, a reverse experiment was made by subjecting caterpillars of the Naples form to very low temperature in rearing. The result was that none was as light colored as the ordinary German form. From these results Weismann concluded that German and Naples forms are constitutionally (genetically) distinct. Weismann (1892, p.401) said, 'A somatogenic character is not inherited in this case, but the modifying influence--temperature--affects the primary constituents of the wings in each individual, i.e. a part of the soma--as well as germplasm contained in the germ of animals.' Weismann (1892, p. 405) even went so far so to say, "In many animals and plants influences of temperature and environment may very possibly produce hereditary variations." Thus contrary to the prevailing belief, Weismann was a neo-Lamarckist, as Darwin was (1972, p. v.). In 1904 Weismann again referred to P. phlaeas in the same context, although he also referred to cases in which environmental inpacts such as nutrition and climatic factors have not affected the germ plasm (e.g., alpine plants, plant galls). In fact, Weismann's theory as a whole was not inconsistent with the Baldwin effect (discussed in Chap. 5). However, what happened later was that his general statement of refutation of the inheritance of acquired characters alone was taken seriously, and it provided a strong theoretical basis for the development of neo-Darwinism." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.40-41)

"Neo-Lamarckism is a more complete theory of evolution than neo-Darwinism, in which the proximate process is ignored, and its methods should enable us to understand the evolutionary process more fully. It is perhaps important to point out further that, besides Huxley and Simpson already referred to, four other prominent evolutionists of our time, Dobzhansky, Rensch, Mayr, and De Beer, accepted or approved neo-Lamarckiam." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.45)

"It should be remembered that, in bringing about macroevolutionary changes, new environmental stimuli in the new (stressful) environments must have induced alteration in gene regulation pathways (Proximate process) and natural selection must have cooperated in completing macroevolutionary changes (ultimate process). This cooperation of the proximate and ultimate processes has been pointed out many times in the name of adaptive response.... To repeat some examples (for the sake of emphasis): embryonization, which occured as a result of interaction between new environmental stimuli and animals, was adaptive for the animals that migrated from water to land (e.g., plethodontids, pulmonates), because the resultant elimination of vulnerable larval stages (on land) was of definite advantage for survival, and natural selection must have contributed to perfect this proximate process byu eliminating those individuals failing to produce large enough eggs. Similarly, neoteny of Ambystoma spp. has occured at high altitudes in North America by the influence of cold temperature on the endocrine mechanism. The resultant neotenous morph was adaptive locally, and natural selection must have favored the individuals that could remain larval (neotenous). Thus in these instances what we see as "adaptive response" today is the product of intricate cooperation of the animals' genotypic response to new environmental stimuli and natural selection. Furthermore, it should be realized that the adaptive response often becomes genetically fixed (Fig. 5) and results in macroevolutionary changes of structures." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.48)

"It is important to note further that what I call "adaptive response" was apparently the way Mivart (1871) understood evolution." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.48)

"It has now become clear that neo-Lamarckism has always been a reasonable theory, and it has stood the test of time for more than a century. Once some misunderstandings and inhibitions are removed, the theory can be regarded as a more complete theory (than neo-Darwinsim) in that it analyses the evolutionary process in terms of both the proximate and ultimate mechanisms, and in that it is especially suited for analyzing the origin of macroevolutionary change. Through the analysis of the proximate process we come to know the cause of variation and the presumed initial stage of evolution of the structures upon which natural selection has worked. In traditional neo-Darwinism natural selection is considered to be involved throughout the whole evolutionary process (of structures), which is indeed untrue, as Mivart (1871) already knew. In practice obvious cases of overextension of the theory of natural selection, which actually results from neglect of the proximate process, have oftne been criticized in terms of their falsifiability. Yet the critics have never offered a solution for this dilemma. Indeed, evolutionary biology has been in a state of constipation caused by the neo-Darwinian constraint that inhibits exploration of the proximate process of evolution. It should now be realized that such a worry will be over once we accept the neo-Lamarckian approach. The application of the neo-Lamarkian analysis appears to resolve some outstanding problems and riddles in evolutionary biology. For instance, the problem of "inheritance of acquired characters" is now understood as the result of accumulation of genocopies. The age-old riddle of "Which came first, the chicken or the egg?" can now be answered from the evolutionary viewpoint (Sect. 3B2). "Adaptive response" nowmust be restored as a fundamental evolutionary concept, though it has been neglected. All phenomena of abnormal metamorphosis (halmatomorphosis, neoteny, caenogenesis) resulting in macroevolutionary structural changes are now attributed primarily to environmentally induced alteration in the response of the genotype (alteration in gene regulation) during the proximate process. The study of the Badwin effect as special cases of genetic assimulation must be encouraged. It should be realized that all the above problems can be more clearly understood by inquiring into the hormonal mediation that becomes involved. Indeed, the environmentally induced hormonal intervention controlling gene action was the mechanism that was unknown to the nineteenth century neo-Lamarckists, and the lack of knowledge of such a mechanism might have hindered the acceptance of neo-Lamarckism." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.53)

"In discussing the evolutionary process constant use of the term neo-Lamarckism has bothered my conscience, because the contribution of Darwin, that is, the part of the evolutionary process played by natural selection, becomes hidden under this term. More properly, therefore, neo-Lamarckism must be called "Lamarck-Darwinism." This again, however, does not convey properly what this work advocated, for both Lamarck and Darwin wrote of many things in addition to the environmental effect on development and evolution (Lamarck) and natural selection (Darwin). To resolve this dilemma and to provide a more precise meaning of what this work emphasizes, a new term, "pan environmentalism," is proposed. In pan-environmentalism, environment consists of both morphogenetic and selective factors. It is envisaged that the former induces, by response of the genotype, variation upon which the selective factor(s) works (Fig 5). It follows, then, that there will be appreciable evolution with environmental changes. (Conversely, there will be no appreciable evolution without environmental change.) Neo-Darwinism may be retained as a method of analysis of the evolutionary process where the effect of environmental change on development is minor or negligible." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.45)

"Numerous enviromental factors (carbon dioxide tension, population density, temperature, photoperiod, etc.) have been known to influence the growth and sexuality in hydras (Loomis 1957, 1959, Burnett and Diehl 1964, Burnett 1968, Schulz and Lesh 1970, Gurkewitz et al. 1980, Shoskak 1981). These environmental factors affect the morphogenesis of hydras by influencing the activity of the four kinds of morphogens or hormones, which consist of an activator and inhibitor for head formation and an activator and inhibitor for foot formation; the two activators are peptides with molecular weights of about 1000, and the inhibitors are peptides with modecular weights (500) (Schaller et al. 1979). It is important to point out further that mutants known to affect morphogenesis of hydras (non-budding, aberrant, maxi, mini, discovered by Schaler el al. 1977a,b,c) control the quantity of localized hormones (morphogens). (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.62)

"It is probably safe to assume that such an endocrine mechanism for vitellogenesis occurs widely within the Mollusca. Presumably, in producing large yolk-rich eggs, altered environmental stimuli (lower salinites, lower temperatures, extreme annual photoperiodic cycle, environmental stimuli on land) have induced the alternation in functioning of the above endocrine mechanism for vitellogenesis (which responds to environmental signals). Hormone(s) concerned with somatic growth and metamorphosis has not been found for the Mollusca, although the existence of such hormone(s) is highly probable. Acceleration in development in enlarged eggs must have involved the alteration in the action of morphogenetic hormones." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.117)

[locusts] "Effects of other environmental factors (such as photoperiod, temperature, nutrition, and humidity) on development and reproduction have been known for a long time (see Mordue et al. 1970, Joly 1972 for reviews of earlier works). Among more recent studies, Albrecht and Lauga (1978) showed that increasing day length and high temperatures have a solitarizing effect in gregarious Schistocerca. Some studies show the direct effect of environmental changes on hormonal activity, which results in the change in development and reproduction. (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.172)

Some species of crickets (Gryllus, Acheta, Gryllodes, etc.) are known to be polymorphic with respect to wing development, both in laboratory experiments and in field observations. Effects of various environmental factors (photoperiod, temperature, population density, food) on wing development have been studied in laboratories. In Gryllodes sigillatus from Canada, which is micropterous in both sexes, Ghouri and McFarlane (1958) found tht the optimal level (for both growth and wing development) is 20% protein in the food. They failed to see the group effect in producing macroptery (probably because of the unfavorable photoperiod). Mathad and McFarlane (1968) found that the exposure of the nymphs to a 10-hr photoperiod suppresses wing development completely, and that a 14-hr exposure is optimal for wing development. They found further that adults exposed to an 14-hr photoperiod produce more winged progeny than adults exposed to an 18-hr photoperiod. In the same species from Japan, however, Arai (1978) found tht the macropterous form appears even in the isolated condition. He showed various photoperiodic effects at different temperatures. Thus all three factors, photoperiod, temperature, and population density influence wing development in Gryllodes sigillatus. Of these, photoperiod appears to be more important factor than the others." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.177-8)

"Wing reduction has occured in many groups of Heteroptera, and aptery sometimes became permanent. Southwood (1961), after reviewing the facts then available, postulated a theory of the cold temperature-hormonally induced wing reduction in Heteroptera. The theory was based on the endocrinological facts brought forward in the studies of Phodnius by Wigglesworth (1952, 1954). Its essential points were as follows: the short-wingedness is a juvenile character and is likely to arise through the change inthe concentration of JH that is necessary for the long-winged adults. This change could be brought about either one of the two ways" (1) by excessive influence of JH leading to juvenile characters --- metathetely, or (2) by depression of the influence of JH (and thus the loss of the prothoracic glands and thus failure of molting hormone), leading to adult characters in the nymph (larva of Southwood) --- prothetely. This view of regarding wing reduction as an aspect either of metathetely (leading to metagenetic neoteny) or of prothetely (leading to progenetic neoteny) has turned out to serve as a very important concept that can be extended to explain two kinds of neoteny in insects as well as in other animals." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.194)

"All of these facts lead me to suspect that embryonization of early larval stages has occured in the enormously enlarged egg (acceleraton), and the result appears to be contracted larval stage and the small adult (progenetic neoteny), as in some dwarf animals producing large eggs." (Matsuda, Ryuichi (1987) Animal Evolution in Changing Environments, with Special Reference to Abnormal Metamorphosis. N.Y.: Wiley Press pp.210)

"Another kind of turnover-pulse hypothesis has been articulated by Calvin (1991), who emphasizes the possible role of glaciation cycles in facilitating human evolution. Until recently, Pleistocene glaciation was assumed to affect primarily upper latitudes, but recent paleoclimatology studies have suggested that glaciation had substantial effects on the temperature, rainfall, seasonality, and patchiness of resources in the equatorial zones of Africa (Foley, 1987). So there is a coincidence between the onset of glaciation cycles and the onset of encephalization inour lineage." (Miller, Geoffrey F. (1994) Evolution of the human brain through runaway sexual selection: the mind as a protean courtship device. unpublished theses. pp. 28)

"...the discoveries of molecular genetics in the course of the last decade have finally demolished Weismann's doctrine of the 'unalterablility of the germ-tract' and modified its modern version, the 'central dogma'. Taken in conjuction with the criticisms discussed earlier on, they may signal the beginning of the end of neo-Darwinism as represented in comtemporary textbooks. Darwinian selection no doubt plays a part in the evolutionary process, but only a subordinate part (comparable to the action of the selective weedkiller) and there is a growing realization that there must be other principles and forces at work on the vast canvas of evolutionary phenomena. In other words, the evidence indicates that evolution is the combined result of a whole range of causative factors - some known, others dimly guessed, yet others so far completely unknown." (Koestler, A. (1978) Janus: A Summing Up: London, Hutchinson)

The connection betwen neoteny and human sexuality and the inverse relationship between aggressive display and sperm production

"But although silver foxes had been bred in captivity since 1892 on fur farms, and although some selective breeding for traits such as fertility and high fur quality had been practiced, the animals were not domesticated in any strict sense. They retained all of the essential characteristics of their wild counterparts. They molted and came into heat ina strict seasonal cycle, as in the wild; their behavior toward humans was no different than that of a wild fox raised in captivity. So Belyaev decided to try an experiment. He would rigorously select animalsapplying but a single criterion: Those animals that shoed consistently tame behavior toward humans would be kept; those that did not would be eliminated from the breeding program. Within five generations changes were already apparent. By 1979, twenty years into the experiment, the results were astonishing. His tame-selected foxes were not just tame; they acted for all the world like domestic dogs. They approached familiar persons and licked their hands and faces. They barked like dogs. They even sought the attention of strangers by whinning and wagging their tails. Their annual molting cycle was disrupted, and the females began to come into heat twice a year, like dogs, and unlike both foxes and wolves. they also developed some physical characteristics of young foxes, such as drooping ears, and some of the variations in traits seen in other domesticated animals, such as piebald coat coloration." (Budiansky 1992: 96, The Covenant of the Wild)

"The time of conception is another nongenetic random variable that may well significantly influence laterality. Seasonal effects have often been considered narrowly. The fact that schizophrenics are more likely to be born in January than July, a finding documented repeatedly has often been interpreted as a result of increased susceptibility of newborn infants to virus infection in the winter. There are many other possibilities, however. Consider, for example, changes in sex hormones with day length. The pineal gland, activated in the dark months, tends to suppress gonadal hormonal production. When it is suppressed, during periods of long days, sex hormones rise. We have already alluded to Badian's (1983) report of a higher rate of nonrighthandedness in males conceived from December through May (days being shortest on December 21 and increasing in length for the following six months). A pineal role in laterality has no direct experimental support, but it certainly deserves study."
(Geschwind & Galaburda 1987: 135-6, Cerebral Lateralization)

"Environmental factors can be an important source of nongenetic influences on laterality. Since the effect of a gene is to play a role in some form of chemical reaction, it is not surprising that genetic determination is not absolute. Every chemical reaction can be modified by alterations in pressure, temperature, pH, light, the presence of other substances, the availability of chemical precursors, and the rate at which products are removed. With growing sophistication of molecular genetics, it has become increasingly clear that nongenetic effects can play a powerful role; methylation, for example, has been shown to suppress expression of many genes. We will now consider some of the random effects that might modify lateralization. One implication of our hypothesis is that even if the genetic endowment of any particular fetus were known precisely, it would not be possible to make predictions concerning the distribution in a population basis. One of the reasons for this relative freedom from genetic determination is that if hormones do play a role in determining laterality, then the effects of testosterone or related substances on the developing brain will be modified by factors not under the control of the fetal genes. Androgens are produced not only by fetal testes and the placenta but also by the maternal ovaries, adrenals, and nonglandular tissues. The fetus can be influenced by the actions of many of the unshared maternal genes. It is reasonable to expect that if a fertilized ovum were transplanted into the uterus of an unrelated female, the final pattern of the brain would be quite different, because the brain would develop in an environment of hormones and other substances that would certainly differ in many respects. It might therefore be reasonable to take a different approach than usual to the genetics of many condiditons. One should perhaps consider, not the genes carried by the offspring alone, but rather the genes of that organism existing or active only for the nine months of pregnancy; in other words, one should consider the mother and the fetus as a unit. This unit contains three groups of different genes: one paternal set present in the fetus, one maternal set present in the mother, and another maternal set present both in the mother and in the fetus. The situation is even more complex when dizygotic twins are involved, since the maternal-fetal unit will contain another group of paternal genes. The effects of substances produced by the mother will, however, be diminished by the capacity of the placenta to act as a barrier to some maternal hormones. The fetus is protected to a great extent form maternal testosterone, which is converted to estradiol by placental aromatase. Dihydrotestosterone, which is not aromatized and therefore crosses the placenta, is, however, usually present in the mother at much lower levels than testosterone. The protection from maternal testosterone is not complete, since offspring do show signs of masculinization when mothers are exposed to this hormone. In addition, progesterone administered to the mother may masculinize female fetuses. It is clear that the placental barrier is far from complete. Furthermore, it is likely that there are individual variations in the aromatizing capacity of the pla
centa. It is conceivable that some maternal genes not shared by the offspring have greater effects on females fetuses. Thus, the testosterone to which female fetuses are exposed comes predominantly from maternal tissues, whereas males produce it themselves in high quantities. In the study of Nichols and Chen (1981) sex hormones given to mothers were associated with a higher rate of hyperactivity in female offspring than in males." (Geschwind & Galaburda 1987: 133-134, Cerebral Lateralization)

"The time of conception is another nongenetic random variable that may well significantly influence laterality. Seasonal effects have often been considered narrowly. The fact that schizophrenics are more likely to be born in January than July, a finding documented repeatedly has often been interpreted as a result of increased susceptibility of newborn infants to virus infection in the winter. There are many other possibilities, however. Consider, for example, changes in sex hormones with day length. The pineal gland, activated in the dark months, tends to suppress gonadal hormonal production. When it is suppressed, during periods of long days, sex hormones rise. We have already alluded to Badian's (1983) report of a higher rate of nonrighthandedness in males conceived from December through May (days being shortest on December 21 and increasing in length for the following six months). A pineal role in laterality has no direct experimental support, but it certainly deserves study."
(Geschwind & Galaburda 1987: 135-6, Cerebral Lateralization)

"Accelterated development was found in brains of infants whose mothers were given adrenal steroids in pregnancy (Barmada and Moossy 1981). (Geschwind & Galaburda 1987: 180, Cerebral Lateralization)

"The possibility of metabolic differences between those with anomalous dominance and those with standard dominance derives some support from Coursey, Buchsbaum, and Murphy's (1979) finding that an excessively high percentage of males with low platelet MAO levels are lefthanded." (Geschwind & Galaburda 1987: 216, Cerebral Lateralization)

"The signigicance of this came out of these first NIMH studies, when it was discovered that one of the primary regulators of the amount of serotonin in the brain was the amount of light a person was exposed to. People with Seasonal Affective Disorder (SAD), a form of wintertime depression, had lower serotonin levels during the winter months. Their SAD went away, however, when they were exposed to several thousand lux of light every day through the use of "light boxes" that they would sit and look at. Other research has implicated light in levels of dopamine, another important neurotransmitter which has been implicated in ADD." (Hartmann 1996: 187, Beyond ADD)

"Regardless of the complexity of sex differences in human neural asymmetries, the close relationship between gonadal hormones and neural sex differences in other species suggests the possibility of a relationship to the early hormonal environment. Data from at least one study support this suggestion. In this study, women who had been exposed to DES for at least 5 months of prenatal life, including the presumed critical period for hormonal influences on human development, were found to differ from their unexposed sisters in performance on a verbal dichotic task composed of consonant-vowel stimuli. As would be expected, given the masculinizing influences of DES on neural and behavioral development in other species (see the first section of this chapter for a discussion of some of these influences), their pattern of performance was shifted in the masculine direction (see Figure 2). Like men in a pilot study using the same dichotic task, as well as in other studies using similar tasks, they showed an enhanced right-ear advantage and a strong negative correlation between right- and left-ear scores. These differences did not seem to be due to other factors, such as age, birth order, pregancy complications, socioeconomic status, or educational background. Nor were they secondary to changes in personality, intelligence, or physical development, all of which were similar in the DES exposed women and their unexposed sisters. Rather, the most likely explanation for the differences between the two groups in performance on the dichotic task seemed to be the early hormonal environment." (Benson and Zaidel (Hines and Gorski)1985: 80, The Dual Brain)

"A study reporting a relationship between rate of maturation and neural asymmetry might also be interpreted to suggest hormonal influences. This study reported that performance on a verbal dichotic task varied with maturation rate, as reflected in the development of secondary sexual characteristics. Individuals with late maturation (i.e., reduced secondary sexual characteristics for their age) showed an enhanced right-ear advantage in comparison to those with early maturation (i.e., enhanced secondary sexual characteristics for their age). This relationship is often interpreted to suggest causal relationship between the rate of neural maturation and the degree of lateralization. Because gonadal hormones determine the development of secondary sexual characteristics, however, the cause of the enhanced lateralization could be hormones rather than maturation rate. A third relationship that has been interpreted to suggest gonadal hormone influences on the development of neural asymmetries involves left-handedness, immune disease, migraine, and developmental learning disorders. Geschwind and Behan reported that individuals who were stongly left-handed were more likely to exhibit learning disorders (i.e., dyslexia and stuttering) and certain immune diseases (e.g., thyroid and bowel disorders) than those who were strongly right-handed. They also found that individuals who suffered from migraine or myasthenia gravis were more likely than controls to show some preference for use of the left hand. The authors suggest that the cause of the associations observed might be elevated testosterone levels during development. They speculate that high levels of testosterone produce left-handedness and learning disorders by delaying the growth of the left hemisphere in utero, and influence the maturation of the immune system by suppressing the thymus. Of interest to the present discussion is the suggestion that prenatal elevations in testosterone may be related to the development of left-handedness. This possibility is consistent with the increased incidence of left-handedness among men and boys (see discussion below), but not with the evidence available from studies of individuals exposed to high levels of testosterone or other gonadal hormones in utero. These individuals have been reported to show normal incidence of left-handedness. Larger samples may be needed to detect a hormonal influence on handedness, however, since the associations reported by Geschwind and Behan were small. Alternatively, the relationship among left-handedness, learning disorders, and immune diseases could be caused by other factors (e.g., the genome or developmental trauma). (Benson and Zaidel (Hines and Gorski) 1985: 82, The Dual Brain)

"The fundamental pattern of the brain thus appears to be asymmetrical, with the same pattern of asymmetries found in most adults. There are, however, influences in pregnancy tht tend to diminish the extent of left-sided predominance, at least in the regions involved in handedness and language, and thus secondarily to result in larger regions on the right side. As noted earlier, our hypothesis is that some factor related to male sex, perhaps testosterone or some closely related factor, is the most likely candidate. The net effect of these intrauterine influences is to produce a shift from left predominance to symmetry, and in a smaller number of cases to modest right predominance." (Geschwind & Galaburda 1987: 46, Cerebral Lateralization)

"Hier and Crowley (1982) found that congenitally hypogonadal males, who lack testosterone, typically had superior verbal scores and low scores on tests of spatial function, unlike normal males." (Geschwind & Galaburda 1987: 97, Cerebral Lateralization)

"Although the majority of human females exposed to DES [diethylstilbestrol] resemble other women on the average, certain features are more common in this population; for example, DES has been shown to have masculinizing effects of the structure of the genitalia (Bongiovanni, Di George, and Grumbach 1959). DES daughters have been reported to have an elevated rate of menstrual difficulties, infertility, and problems during pregnancy, including premature and preterm births, prolonged labor, and higher rate of children with congenital anomalies (Herbst et al. 1980), though agreement on this is not universal. However, it should be noted that the mothers of these women also had some of these problems, which probably constituted the reason for their taking DES. It will be necessary to study a large number of DES daughters in order to ascertain whether they show an increased rate of the changes in the cortex that we have hypothesized would result from high intrauterine testosterone exposure. One might expect DES daughters to have an elevated frequency of lefthandedness, learning disabilities, high spatial talents, and the other associations hypothesized to be more frequent with elevated intrauterine male-related effects. Hines (1982) has reviewed the effects of DES. We have carried out a study on a group of 77 adults exposed to DES in utero, and our preliminary results show a markedly elevated rate of non-righthandedness in this population. Nichols and Chen (1981) have found that sex hormones administered to pregnant women lead to an elevated rate of hyperactivity in the offspring, especially the girls." (Geschwind & Galaburda 1987: 111-112, Cerebral Lateralization)

"Even if it is true that a male-related factor modulates intrauterine growth of certain cortical regions, it is obvious that cortical development will be susceptible to many other influences, some closely related to the hormonal effects, some more remote. Ward and Weisz (1980) showed that if a female rat is stressed during the course of pregnancy, the male offspring show demasculinization. After the stress there is a rapid rise of testosterone in male fetuses, followed by a fall to subnormal levels. It is not clear exactly how stress on the mother leads to this sequence of changes. It is interesting to note that the female offspring of such a stressed mother will also tend to have demasculinized male offspring, even though these famales are not themselves stressed in pregnancy. Phenobarbital administered during pregnancy has similar effects, and the male offspring have permanently low testosterone levels.This is an example of a phenomenon already mentioned, whereby chemical effects on the fetus may produce permanent alterations in metabolism. When present in the female they can lead to a mechanism of nongenetic transmission to the following generation. Sherman et al. (1980) have also demonstrated an environmental effect on lateralization. In studies carried out some years earlier it had been shown that brief periods of handling of newborn rats exerted permanent effects on emotionality and on other traits such as resistance to infection. In the more recent experiments it was shown that male rats who had been handled showed a greater right hemisphere lateralization in tests of emotionality and spatial performance than animals who had not been." (Geschwind & Galaburda 1987: 115, Cerebral Lateralization)



"A corollary of our hypothesis is that hormonal effects on the brains of offspring may vary with the time of conception. The activity of the pineal gland changes seasonally with alterations in day length. As a general rule, during the dark winter months the pineal becomes active and suppresses both ovaries and testes, whereas in the summer it is inactive and sex hormone levels are higher. For this reason many animals bear young in the spring, an advantageous situation since temperature and food supplies are more suitable for survival. An example of such seasonal modulation of hormonal effects on the brain is observed in the HVc nucleus of the singing bird (Nottebohm 1981). This description of pineal physiology is, however, somewhat oversimplified. An animal's sensitivity to light may vary through the year. Gonadal hormones may thus become activated in the spring, but as a result of loss of sensitivity to light over th summer hormone levels may diminish as fall approaches. Despite these facts, day length is a powerful influence. Thus, steers increase their weight more rapidly in the winter when artificial light is supplied to lengthen the day. This light-enhanced growth of muscle mass does not take place if the bull is castrated, suggesting that the effect of light is mediated through a rise in testosterone effect (Tucker and Ringer 1982).....If pineal effects on sex hormone levels are important, then the birth months of lefthanders, and of those with learning disorders, might not be uniform throughout the year, since fetuses conceived at different seasons might be subjected to very different hormonal environments. These effects should differ in the Northern and Southern Hemispheres and at the equator, although other factors, such as variations in the ethnic composition of populations, would also have to be considered. Data are still very sparse. Badian (1983) found that in males born in each of the six months beginning in September, the rate of nonrighthandedness was higher than that found in any of the other six months, but no clear trend was observed for female births." (Geschwind & Galaburda 1987: 116-7, Cerebral Lateralization)

"Diethylstilbestrol (DES) was shown years ago to produce masculinizing effects on female offspring (Bongiovanni, Di George, and Grumbach 1959). The reasons for this have become clear in recent years with the discovery that testosterone masculizes the rat brain. Estradiol within certain nuclei of the hypothalamus is actually the final cause of this masculinization, but these nuclei are protected from the circulating estradiol that, in the rat, is bound to alphaafetoprotein (AFP). Testosteone is not bound by AFP and thus reaches these nuclei, where it is converted to estradiol by aromatase. DES also masculinizes; although it is an estrogen, it is not bound by AFP and thus reaches the brain sites within which it acts like estradiol to masculinize. It is not surprising that DES-exposed daughters have a high rate of gynecological difficulties and infertility and an elevated rate of children with congenital anomalies, all of which may reflect this masculinizing effect. We have seen several DES daughters with features such as facial hirsutism, acne, and unusual sensitivity to the masculinizing effects of phenytoin. A high rate of anomalous dominance might be expected in this group, and our preliminary findings of 77 DES daughters support this hypothesis. In psychological studies of a small sample of DES children Hines (1982) found some results that she interpreted as consistent with a masculinizing effect on the brain, but other data summarized in her review differ; the numbers were again small, however. Further study on larger numbers is obviously warranted." (Geschwind & Galaburda 1987: 172-3, Cerebral Lateralization

"In recent years more and more evidence has been found that testosterone affects not just ornaments and bodies but also brains. Testosterone is an ancient chemical, found in much the same form throughout the vertebrates. Its concentration determines aggressiveness so exactly that in birds with reversed sex roles, such as phalaropes and in female-dominated hyena clans, it is the females that have higher levels of testosterone in the blood. Testosterone masculinizes the body; without it, the body remains female, whatever its genes. It also masculinizes the brain. Among birds it is usually only the male that sings. A zebra finch will not sing unless there is suffecient testosterone in its blood. With testosterone, the special song-producing part of its brain grows larger and the bird begins to sing. Even a female zebra finch will sing as long as she has been exposed to testosterone early in life and as an adult. In other words, testosterone primes the young zebra finch's brain to be responsive later in life to testosterone again and so develop the tendency to sing. Insofar as a zebra finch can be said to have a mind, the hormone is a mind-altering drug. Much the same applies to human beings. Here the evidence comes from a series of natural and unnatural experiments. Nature has left some men and women with abnormal hormonal doses, and in the 1950's doctors changed the hormonal conditions of some wombs by injecting some pregnant women with certain hormones. Women with a condition known as Turner's syndrome (they are born without ovaries) have even less testosterone in their blood than do women who have ovaries. (Ovaries produce some testosterone, though not as much as testicles do.) They are exaggeratedly feminine in their behavior, with typically a special interest in babies, clothes, housekeeping, and romantic stories. Men with less than usual testosterone in their blood as adults---eunuchs, for example---are noted for their femininity of appearance and attitude. Men exposed to less than usual testosterone as embryos---for example, the sons of diabetic women who took female hormones during pregnancy---are shy, unassertive, and effeminate. Men with too much testosterone are pugnacious. Women whose mothers were injected with progesterone in the 1950's (to avert miscarriage) later described themselves as having been tomboys when young; progesterone is not unlike testosterone in its effects. Girls who were born with an unusual condition called either adrenogenital syndrome or congenital adrenal hyperplasia are equally tomboyish. This order causes the adrenal gland, near the kidney, to produce a hormone that acts like testosterone instead of cortisol, its usual product. Somewhat like in the zebra finches, there is two periods when testosterone levels rise in male children: in the womb, from about six weeks after conception, and at puberty. As Anne Moir and David Jessel put it in a recent book, Brain Sex, the first pulse of hormone exposes the photographic negative; the second develops it. This is a crucial difference fro the way the hormone affects the body. The body is masculinized by testosterone from the testicles at puberty, whatever its womb experience. But not the mind. The mind is immune to testosterone unless it was exposed to a sufficient concentration (relative to female hormones) in the womb. It would be easy to engineer a society with no sex difference in attitude between men and women. Inject all pregnant women with the right dose of hormones, and the result would be men and women with normal bodies but identical feminine brains. War, rape, boxing, car racing, pronography, and hambergers and beer would soon be distant memories. A feminist paradise would have arrived. (Ridley 1993: 254-6, The Red Queen) [This quote suggests that feminity as suggested by Turner's syndrome, is a default position. It takes testosterone to establish male features. Could there be an automatic neotenous direction of evolution in the event that testerone decreases?]


"When the day is artificially lengthened in the winter, Holstein cows increase their milk production and steers gain weight faster (though not when castrated, which strongly suggests that the testes play a role). It is likely that the pineal gland exerts a major control in this regard. During the dark months it is active and suppresses the gonads, but during the longer days it becomes inactive and the production of gonadal hormones therefore rises. This mechanism clearly makes it more likely that animals will be born in the spring, a useful adaptation since conditions of food and climate are far more favorable for survival. Sensitivity to long days appears to diminish over time, so that it is less marked by the end of the summer (Reiter 1980). Male rats raised in constant darkness show diminished sexual activity, but this is reversed by pinealectomy (Baum 1968), presumably because of the removal of pineal inhibition. The yearly cycle is not merely attended by changes in the other metabolic alterations. The hyperglycemic response to epinephrine in humans is higher in the winter than in the summer (Altschule and Siegel 1951). The goldenhamster shows greater thermogenesis in response to a cold stimulus in midwinter than in midsummer (Pohl 1965). Though the yearly metabolic cycle can probably not be attributed entirely to activity of the pineal, it must play a major role. We have already noted that cyclic alteration in the production of sex hormones is likely to be important in many ways; for example, it might affect the percentage of children with anomalous dominance born at different seasons. Children conceived in March or April will spend most of their first months in utero at a time when hormones are high. Children conceived six months later, in September or October, will tend to spend their early period in utero under much lower hormonal influences. Obviously, different quarters of the year will give rise to different patterns, but adequate information regarding the outcomes is still lacking. Badian (1983) found that nonrighthandedness was twice as common among boys born in the six months beginning in September (and thus conceived from December through May) than among those born in the following six months. Furthermore the number of nonrighthanders born in each of these months was higher than the number born in every month in the other half year. Similar effects were not found in females. It is intriguing that compared to controls, rabbits raised postnatally in darkness for seven months were fond to show an increase if synaptic contact zones in the medial visual cortex and the motor cortex on the left, but not on the right (Vrensen and deGroot 1974); however, the authors are cautious in their interpretation of this. There are other possible relevant data. It has been confirmed repeatedly that schizophrenics are born predominantly in the first half of the year particularly in the first quarter. They are thus conceived between approimately April 1 and October 1, that it, predominantly during the period in which the days are longer than 12 hours. It is possible that schizophrenia is more common in individuals who have spent the first six months of pregnancy under maximal hormonal influences. Mental defectives are also more likely to be born at the beginning of the year. On the other hand, many extensive studies of the birth months of eminent people have shown that they too tend to be born predominantly early in the year; even more consistently, the rate of such births is low in the midsummer months of July and August (Peterson 1979). (In all of these studies the data have been corrected for the normal yearly pattern of births.)" (Geschwind & Galaburda 1987: 219-20, Cerebral Lateralization

"Even if it is true that a male-related factor modulates intrauterine growth of certain cortical regions, it is obvious that cortical development will be susceptible to many other influences, some closely related to the hormonal effects, some more remote. Ward and Weisz (1980) showed that if a female rat is stressed during the course of pregnancy, the male offspring show demasculinization. After the stress there is a rapid rise of testosterone in male fetuses, followed by a fall to subnormal levels. It is not clear exactly how stress on the mother leads to this sequence of changes. It is interesting to note that the female offspring of such a stressed mother will also tend to have demasculinized male offspring, even though these famales are not themselves stressed in pregnancy. Phenobarbital administered during pregnancy has similar effects, and the male offspring have permanently low testosterone levels.This is an example of a phenomenon already mentioned, whereby chemical effects on the fetus may produce permanent alterations in metabolism. When present in the female they can lead to a mechanism of nongenetic transmission to the following generation. Sherman et al. (1980) have also demonstrated an environmental effect on lateralization. In studies carried out some years earlier it had been shown that brief periods of handling of newborn rats exerted permanent effects on emotionality and on other traits such as resistance to infection. In the more recent experiments it was shown that male rats who had been handled showed a greater right hemisphere lateralization in tests of emotionality and spatial performance than animals who had not been." (Geschwind & Galaburda 1987: 115, Cerebral Lateralization)

"The flounder had both eyes on one side of its head. In some individuals the eyes are on the right; in others they are on the left. The distribution on the two sides varies in the different species that occupy particular geographic niches. Cross-breeding experiments between species have been carried out, with findings that do not fit readily into any standard nuclear genetic model (Policansky 1982). It is possible that variations in the distribution to asymmetry may be deterined to a great extent by differing environmental conditions, such as water temperature or food supply. This possibility gains credence form the fact that sex ratio of the offspring of certain species shifts sharply from female predominant to male predominant depending on temperature or food supply. (Clutton-Brock 1982); Harvey and Slatkin 1982). In several species the sex ratio can vary considerably depending on the hormonal conditions of pregnancy. James (1980a,b) has suggested that this may be true even in humans. It is thus also conceivable that there is a genetic bias toward one form of laterality, which is modified by environmental conditions that alter hormonal milieu or the time of fertilization." (Geschwind & Galaburda 1987: 128-9, Cerebral Lateralization)

"There is another possible but surprising souce of random variation. When rats are subjected to stress during pregnancy, the offspring show altered emotional behavior. In turn, the offspring of affected female offspring show similar changes in emotional behavior despite the fact that their mothers were not exposed to any additional stress. This can readily be explained. The offspring of a mother subjected to stress in pregnancy show permanent alterations in their own behavior and are more susceptible to stress effects than normal animals. Therefore, when the females among them become pregnant, even minor stresses will affect them, and their offspring will in turn be affected---an example of what might loosely be termed a "Lamarckian" effect. There are two mechanisms by which such persistent effects of stress could be mediated. Stress produces structural alterations in the brain of the offspring and/or permanent metabolic alterations. Thus, newborn female rats exposed to testosterone manifest a series of permanent metabolic changes. They are more sensitive in later life to androgens than normal females. They carry sex-linked protein, which is normally found only in males (Michaelson 1981), and certain liver enzymes normally found in females disappear. (Gustaffson et al. 1978). There is thus an increased possibility that the female who was subjected to excessive testosterone effects in utero will herself show an increased tendency to hormonal masculinization. which will in turn affect her offspring. In the rat diethylstilbestrol (DES) has masculinizing effects on brain regions involved in reproduction behavior (MasLusky and Naftolin 1981). In the human DES also appears to have certain masculinizing effects (Bongiovanni, Di George, and Grumback 1959); moreover, as mentioned earlier, DES daughters have been reported in some series to have a higher rate of infertility and difficulties in pregnancy, although this result is controversial (Beral and Colwell 1981). We hypothesize that these women will have a higher proportion of children with anomalous dominance, that is, a higher rate of lefthandedness and of the other associations of atypical laterality. Preliminary studies now under way appear to confirm this. There will be certain limitations on such transgenerational masculinizing effects. Even if the mother is masculinized by endogenous of exogenous hormones, the fetus may be shielded from them to a varying degree by the placenta and by the effects of paternal genes." (Geschwind & Galaburda 1987: 136-7, Cerebral Lateralization)

There will also be nongenetic effects. Thus, when the mother is anomalously dominant, she will often be hormonally anomalus in such a way as to favor the production of children with similar dominance patterns. the anomalous hormonal pattern of the mother may reflect her own genetic pattern, but when the responsible genes are not shared with the fetus, then the effects on the fetus will be independent to a great extent of its own genetic endowment. There will be other cases in which the mother was herself exposed to an anomalous hormonal environment, as a result of her own genetic endowment or as a result of nongenetic effects, for example, hormones controlled by maternal genes that she did not share or exogenous stimuli that altered the hormonal atmosphere, such as sex steroids, other drugs, and even the season of birth." (Geschwind & Galaburda 1987: 177, Cerebral Lateralization)

"Huntington's disease is transmitted as a Mendelian dominant; that is, on the average half of both male and female children are affected, and the disease is passed on equally often by affected fathers and affected mothers. Unexpectedly, however, early onset cases of this disorder are overwhelminly often transmitted by the father (Myers et al. 1983). (Geschwind & Galaburda 1987: 177, Cerebral Lateralization)

"E. J. Quart (personal communication) found an elevated rate of a past history of dyslexia in a group with Reye syndrome. This intriguing possibility deserves further study, especially in view of the evidence that this disorder involves disturbance of mitochondrial (dytoplasmic) enzymes. We have discussed already, and will discuss again, the possible rode of cytoplasmic inheritance as a determinant of laterality, as suggested by Corballis and Morgan (1978). (Geschwind & Galaburda 1987: 198, Cerebral Lateralization)

"The hormonal atmosphere in utero may well permanently alter expression of genes or alter genes themselves (for instance, by methylation). (Geschwind & Galaburda 1987: 218, Cerebral Lateralization)

"When the day is artificially lengthened in the winter, Holstein cows increase their milk production and steers gain weight faster (though not when castrated, which strongly suggests that the testes play a role). It is likely that the pineal gland exerts a major control in this regard. During the dark months it is active and suppresses the gonads, but during the longer days it becomes inactive and the production of gonadal hormones therefore rises. This mechanism clearly makes it more likely that animals will be born in the spring, a useful adaptation since conditions of food and climate are far more favorable for survival. Sensitivity to long days appears to diminish over time, so that it is less marked by the end of the summer (Reiter 1980). Male rats raised in constant darkness show diminished sexual activity, but this is reversed by pinealectomy (Baum 1968), presumably because of the removal of pineal inhibition. The yearly cycle is not merely attended by changes in the other metabolic alterations. The hyperglycemic response to epinephrine in humans is higher in the winter than in the summer (Altschule and Siegel 1951). The goldenhamster shows greater thermogenesis in response to a cold stimulus in midwinter than in midsummer (Pohl 1965). Though the yearly metabolic cycle can probably not be attributed entirely to activity of the pineal, it must play a major role. We have already noted that cyclic alteration in the production of sex hormones is likely to be important in many ways; for example, it might affect the percentage of children with anomalous dominance born at different seasons. Children conceived in March or April will spend most of their first months in utero at a time when hormones are high. Children conceived six months later, in September or October, will tend to spend their early period in utero under much lower hormonal influences. Obviously, different quarters of the year will give rise to different patterns, but adequate information regarding the outcomes is still lacking. Badian (1983) found that nonrighthandedness was twice as common among boys born in the six months beginning in September (and thus conceived from December through May) than among those born in the following six months. Furthermore the number of nonrighthanders born in each of these months was higher than the number born in every month in the other half year. Similar effects were not found in females. It is intriguing that compared to controls, rabbits raised postnatally in darkness for seven months were fond to show an increase if synaptic contact zones in the medial visual cortex and the motor cortex on the left, but not on the right (Vrensen and deGroot 1974); however, the authors are cautious in their interpretation of this. There are other possible relevant data. It has been confirmed repeatedly that schizophrenics are born predominantly in the first half of the year particularly in the first quarter. They are thus conceived between approimately April 1 and October 1, that it, predominantly during the period in which the days are longer than 12 hours. It is possible that schizophrenia is more common in individuals who have spent the first six months of pregnancy under maximal hormonal influences. Mental defectives are also more likely to be born at the beginning of the year. On the other hand, many extensive studies of the birth months of eminent people have shown that they too tend to be born predominantly early in the year; even more consistently, the rate of such births is low in the midsummer months of July and August (Peterson 1979). (In all of these studies the data have been corrected for the normal yearly pattern of births.)" (Geschwind & Galaburda 1987: 219-20, Cerebral Lateralization)

"Further work showed that for most genes, corrections would be made---splicing out the nonsense---after an RNA copy is made of a DNA gene. Even with "interrupted" DNA, an edited and corrected message in RNA could be used by the cell's machinery to make the correct protein. Even more surprisingly, for antibody genes the DNA itself can also be spliced. In other words, DNA that is inherited can be altered. Amazing!" (Behe 1996: 127, Darwin's Black Box.)

"Fed on identical diets, two genetically different men will not grow to the same height. Fed on different diets, two identical twins will grow to different heights. Nature is the length of the rectangle, nurture the width. There can be no rectangle without both. The genes for height are really only genes for responding to diet by growing." (Ridley 1993: 265, The Red Queen)

"My own expectation is that when the almost totally unknown realm of processes whereby DNA determines embryology is studied, it will be found that DNA mentions nothing but relations." (Bateson 1979: 174, Mind and Nature)

"That genetic change can at least partly avoid the price of imposing rigidity on the system by being delayed until it is probable that the circumstance which was coped with by the soma at a reversible level is indeed permanent and by acting only indirectly on the phenotypic variable. The genetic change presumably shifts only the bias or setting (see Glossary, "Logical Type") of the homeostatic control of the phenotypic variable." (Bateson 1979: 175, Mind and Nature)

"It is of primary importance to note that insofar as embryos are protected in eggs or in the mother's body, the external environment will not have a strong selective effect on genetic novelties until epigenesis has proceeded through many steps. In the past and still continuing into the present, extenal natural selection has favored those changes that protect the embryo and juvenile from external dangers. The result has been an increasing separation between the two stochastic systems." (Bateson 1979: 196, Mind and Nature)

"But, to borrow an old metaphor from Francis Galton (see Essay 27 for a full explication), suppose that organisms are polyhedrons rather than billiard balls---and that they can only move by flipping from one side (on which they now rest) to an adjacent facet. They may need a push from natural selection to move at all, but internal ilimitations and possiblities set the direction of possible change....But these same channels, properly interpreted as strong biases in variation rather than one-way streets of change, would give us a much richer view of evolution as a subtle balance between constraints of history and reshapings by natural selection." (Gould 1993: 368-9, Eight Little Piggies)

"There is indeed some evidence that females who are high and males who are low in the male sex hormone (androgen) score higher on spatial ability tests (see McGee, 1979). While this would support the idea of the need for an optimal (i.e. intermediate) level of hormones...." (Bradshaw & Nettleton 1983: 218, Human Cerebral Asymmetry)

"Left-handers are not a homogenous group with respect to patterns of lateralization, however. Clinical and experimental data reveal an important difference between familial and nonfamilial left-handers. Nonfamilial left-handers appear to be more like male right-handers; the incidence of aphasia is consistently associated with lesions to only one side of the brain, the left hemisphere. Initial apahsia can be more severe and recovery from aphasia slow and limited as well. But familial left-handers and those right-handers with left-handedness in the family appear to have greater bilateral control of language and better prognosis for recovery. In experimental data as well, cerebral ambilaterality is more associated with familial sisistrality, while nonfamilial left-handers show more consistent unilateral left-sided dominance for language." (Benson and Zaidel (Curtis) 1985: 105, The Dual Brain)

"Furthermore, Annett (1970) has stressed that handedness, like height, is a continuous variable. In studying the associations of handedness, the appropriate question is therefore not, "Is lefthandedness more common in condition A than is the general population?" but rather, "Is the distribution of handedness scores different in condition A from that in the general population?" (Geschwind & Galaburda 1987: 75, Cerebral Lateralization)

"Other features of the familial distributions present problems for any genetic theory, for example, a trend for lefthanded mothers more than lefthanded fathers to have lefthanded children." (Geschwind & Galaburda 1987: 128, Cerebral Lateralization) [Question: do the females come from more familial lefthanded families? Since more males than females are lefthanded, is that a result of higher testosterone levels over and above present male levels causing a shift further back to circle culture laterialization?][Is the fact above indicative of Lemarckian influences more responsive to in vitro environment of left handed mother?] [about more males being pathologically lefthanded.]

"Thus, too studies (Churchill, Igna, and Seirf 1962; Grapin and Perpere 1968) have shown a strong relationship between birth position and later handedness, with ROA [right occiput anterior] infants having a much higher later frequency of lefthandedness than LOA infants." (Geschwind & Galaburda 1987: 226, Cerebral Lateralization)

"A resolution to this issue if offered by Annett and Turner (1974), who showed that when a large random sample is classified for handedness, and the resultant groups compared, for example, on reading scores, no significant differences emerge. However, if one then looks only at those with very low scores, more sinistrality is pathological. Confirmation is offred by Bishop (1980), who took an unselected group of preadolescents, tested them on intelligence, reading ability, and manual dexterity, then chose a target group not on the basis of sinistrality but on the basis of very poor performance by the nonpreferrred hand (i.e., usually but not always the left hand). These subjects were impaired in intelligence and reading, and sinistrals in the target group had a lower incidence of famailial sinistrality than other sinistrals, who were not as impaired in performance on the nonpreferred (right) hand. These target-group (nonfamilial) sinistrals were presumably pathological left-handers, perhaps with mild unilateral brain abnormality leading to a depression of contralateral performance, clumsiness, poor cognitive performance, and sinistrality. In two of our own studies (J.L. Bradshaw et al., 1981a; J.L. Bradshaw & Taylor, 1979), we found in a sample of university students that nonfamilial sinistrals performed significantly more poorly on tachistoscopically presented verbal tasks (naming word-non-word decisions) than either familial sinistrals or dextrals. We must, however, conclude that across the whole population, excluding university students and clinical samples, which between them seem to have attracted an undue degree of research attention to date, there is comparatively little evidence of any obvious deficit in sinistral functioning. Apart from birth stress hypothesis per se, it is also possible that a language invasion into right hemisphere processing space (which, to judge by their possibly reduced asymmetries might seem to be more common among both sinistrals and females), might make sinistrals inferior to dextrals on nonverbal tasks and possibly superior on verbal abilities (J. Levy, 1969). (Such a double dose of language, apart form boosting the faculty, could also account for the reportedly higher incidence of stuttering among sinistrals because of hemispheric competition for the unpaired articulators.) Females are superior to males verbally, and inferior in nonverbal tasks (see Chapter Eleven). Some empirical support is claimed for the Levy hypothesis, though others have found no differences between handedness groups (for detailed reviews, see Carter-Saltzman, 1979; Swanson, Kinsbourne, & Horn, 1980). Since those reviews were published, O. Johnson & Harley (1980) found that strongly sinistral university students were better on verbal and worse on spatial tasks than strong dextrals. J.L. Bradshaw et al. (1981a) found evidence of a significant nonverbal intelligence deficit among sinistral university students, particularly familial, with parental handedness being a better predictor of performance than sibling handedness. Exactly comparable findings in a dichaptic shape discrimination task were reported by Nilsson et al. (1980): very poor performance by familial sinistrals, especially when the relationship was one of parent rather than sibling sinistrality. Such findings suggest that sinistrals have a somewhat different cognitive organization form dextrals, and that both environmental factors (inferior nonfamilial sinistral processing on speeded tasks) and gentic factors (inferior familial sinistral processing on unspeeded intelligence tests) may operate in sinistrality." (Bradshaw & Nettleton 1983: 208-9, Human Cerebral

"Familial sinistrality, and especially parental handedness, is an important determinant of performance. Nonfamilial sinistrals appear to be the least lateralized or the most likely to demonstrate abnormal laterization. .... A number of genetic models have been proposed, but all so far have foundered on the twin and sibling data." (Bradshaw & Nettleton 1983: 265, Human Cerebral Asymmetry) [note that this seems to contradict another study]

"Boys with an extra X chromosome (XXY instead of the normal XY) are much more verbal than other boys. Girls with Turner's syndrome (no ovaries) are even worse at spatial tasks than other girls but just as good at verbal ones. Girls who were exposed to male hormones in the womb are better at spatial tasks. Boys who were exposed to female hormones are worse at spatial tasks." (Ridley 1993: 257, The Red Queen)

"A disproportionate number of sinistrals are found among a number of clinical populations, including mental retardates, epileptics, stutterers, and sufferers from dysarthria, cerebral palsy, developmental aphasia, and dyslexia (for reviews, see Carter-Saltzman, 1979; Hicks & Kinsbourne, 1978; Satz, Baymur, & Van der Vlugt, 1979; Springer & Searleman, 1980). There are even said to be more nondextrals among smokers (harburg, Feldstein, & Papsdorf, 1978) and criminals (Andrew, 1978)! Sinistrality is also said to be more common among the first- or late-born, those born of very young or old mothers, twins, males (who are larger at birth), and those known to be the products of complicated pregnancies." (Bradshaw & Nettleton 1983: 206-7, Human Cerebral Asymmetry)

"Another example is much more common, but less acknowledged. It is the case of cell differentiation in embryonic development. Once a cell differentiates by responding to an environmental factor (i.e. an inductor substance), it usually passes its characteristics to descendants. (Alberts et. al. 1989, Blau, 1989). Again, after the acquisition of the differentiated character, there is hereditary transmission of it." (Aboitiz, F. (1992) Mechanisms of adaptive evolution. Darwinism and Lamarkism restated. Medical Hypothesis 38(3): pp. 196)

"In the light of modern genetics, it is presently considered that IAC is practically an impossibility. In metazoans, the germline and the genetic material are supposed to be isolated form perturbations coming from the environment. Thus, ontogentic changes may not effect heredity. This assumption seems correct for metazoans, in whom the majority of phenotypic traits is indirectly inherited. However, as already mentioned, interitance of acquired characteristics becomes more plausible in cases in which direct inheritance makes a significant portion of the phenotype, as in cell division. If, for example, a cytoskeletal feature is modified during a cell's ontogeny, this is likely to be inherited by descendants. This point was referred to as the transgenerational dimension of modifications to directly inherited characters." (Aboitiz, F. (1992) Mechanisms of adaptive evolution. Darwinism and Lamarkism restated. Medical Hypothesis 38(3): pp. 196)

"They showed in Table I and II and Figs. 1 and 2. Alcohol patients showed significantly lower testosterone levels than controls. The circadian rhythm was lost after withdrawal but it recurred on third day and progressively increased until full recovery, i.e. the difference between hormone levels at 9 and 21 h increased slowly, reaching nearly the level of signification on day 15. FSH and LH levels were signifantly higher in alcoholics at day 1, and the levels did not vary significantly during the 15 days of the study. A strong relationship existed between levels of FSH and LH (r = 0.56, P< 0.0001), but not between these hormones and testosterone. Prolactin levels significantly increased during the 15 days of observation, being in the interior limit of the normal range at the beginning." (Castilla-Garcia, A., Santolaria-Fernandez, F.J., Gonzalez-Reimers, C.E., Bastita-Lopez, N., Gonzalez-Garcia, C., Jorge-Hernandez, J.A., & Hernandez-Nieto, L. (1987) Alcohol-induced hypogonadism: reversal after ethanol withdrawal. Drug and Alcohol Dependence 20(3): pp. 257)

"In our study, testosterone levels progressively returned to normal along the 15 days of the observation period. In addition, the circadian rhythm, lost at the beginning of the study --- a feature also reported by Gordon (2) and Bertello(22) --- became evident again in the last days of the observation period. These data suggest that hypogonadism seems to be functional in nature. .... Prolactin has been found to be elevated in alcholics after a week of abstinence (13,23), and decreased during the hangover phase, with blockade of the prolactin response to TRH (24). In this connection it has been argued that ethanol withdrawel may lead to an excessive dopaminergic activity, and, as a consequence, to a low release of prolactin (24). In our study, serum prolactin levels-especially those obtained at 21 h --- recovered progressively from the low limit of the normal range observed in the first days to the higher levels at the end of the observational period." (Castilla-Garcia, A., Santolaria-Fernandez, F.J., Gonzalez-Reimers, C.E., Bastita-Lopez, N., Gonzalez-Garcia, C., Jorge-Hernandez, J.A., & Hernandez-Nieto, L. (1987) Alcohol-induced hypogonadism: reversal after ethanol withdrawal. Drug and Alcohol Dependence 20(3): pp. 259)

Chart (book wall) comparing testosterone levels of Chinese, Japanese, African and Whites, all USA. Whites had lowest level of the four, contradicting shift theory expectations. Tests need to be done of ethnic groups within their cultures, that stress increases testosterone levels could heavily influence the figures in this study; the measurements may be noting the relative stress of community ties and socioeconomic status. (Wu, A.H., Whittemore, A.S., Kolonel, L.N., John, E.M., Gallagher, R.P., West, D.W., Hankin, J., Teh, C.Z., Dreon, D.M., & Paffenbarger, R.S.Jr. (1995) Serem androgens and sex hormone-binding globulins in relation to lifestyle factors in older African-American, white, and Asian men in the United States and Canada. Cancer Epidemiology, Biomarkers & Prevention 4(7): pp. 738)

"After adjustment for age, Quetelet's index, and physical activity, levels of testosterone (total, free, and bioavailable) were higher for Asians born in Asia versus for those Asians born in North America, whereas levels of SHBG and DHT were similar in both groups (Table 4). On the other hand, the DHT: testosterone ratio was lower in Asians born in Asia compared to those born in North America. However, none of the differences between Asians born in North America and those born in Asia were statistically significant." (Wu, A.H., Whittemore, A.S., Kolonel, L.N., John, E.M., Gallagher, R.P., West, D.W., Hankin, J., Teh, C.Z., Dreon, D.M., & Paffenbarger, R.S.Jr. (1995) Serem androgens and sex hormone-binding globulins in relation to lifestyle factors in older African-American, white, and Asian men in the United States and Canada. Cancer Epidemiology, Biomarkers & Prevention 4(7): pp. 739)

"Stenuous physical activity may result in lower levels of testosterone (26-28). Smoking may inhibit production of estrogens in women (29) but data on its effect in men are not consistent (30,31). In a study of Caucasian upper middle class men in a California retirement community, smokers showed significantly higher serum estrogen levels than did nonsmokers (30). However, in the Multiple Risk Factor Intervention Trial, serum estrogen levels in men were not associated with total amount of cigarettes smoked (31). Levels of testosterone were higher among men who smoked compared withmen who did not in both studies (30,31). Heavy alcohol use may increase estrone production and, thus, increase conversion of androgens to estrogens (32, 33). The present findings suggest higher total testosterone levels in smokers compared to nonsmokers, but there were few consistent associations between physical activity, alcohol use, and androgen levels. In this study, men with a self-reported history of a benign prostate condition showed a signigicantly higher DHT: testosterone ratio, a marker of 5a-reductase activity (34), compared to men without such history. This observation is compatible withthe hypothesis that the development of benign prostate (35). .... The present study of older men, most of whom were 60 years or older, found lower testosterone levels in whites compared to African-Americans, similar to studies conducted in young college men and Vietnam veterans aged 31-50 years. However, there is little evidence from this and other studies that testosterone levels in Asians residing in Asia or in North America are lower than those in whites. On the other hand, the strongest support for ethnic differences in androgen levels in this study was bsed on the DHT:testosterone ratio, which was highest in African-Americans, intermdiate in whites, and lowest in Asian-Americans, reflecting the prostate cancer incidence rates in these groups." (Wu, A.H., Whittemore, A.S., Kolonel, L.N., John, E.M., Gallagher, R.P., West, D.W., Hankin, J., Teh, C.Z., Dreon, D.M., & Paffenbarger, R.S.Jr. (1995) Serem androgens and sex hormone-binding globulins in relation to lifestyle factors in older African-American, white, and Asian men in the United States and Canada. Cancer Epidemiology, Biomarkers & Prevention 4(7): pp. 739)

"First, both physical and psychological stress have been consistently found to depress T (Mason, 1968; Kreuz et al., 1972; Opstad & Aakvaag, 1982). Second, aggressive or competitive/dominance encounters generally raise T levels, which also reflect the outcome of the encounter: T levels are elevated in winners and relatively repressed in losers (Rose et al., 1975; Elias, 1981). ... Generally, sustained moderate-to-intense exercise appears to elevate T, although data from studies have been equivocal (Kuoppanalmi et al., 1980)." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 450)

"In Western clinical populations, a marked diurnal variation in serum T levels has been reported, with evening values averaging around 30% lower than morning values (Nieschlag, 1974). Baseline values of !Kung hunters show similar diurnal variation." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 452)

"Evolutionary arguments for the importance of hunting in shaping foundations of men's behavior have tended to focus on its aggressive, competitive and stressful components. Its basic nature as a form of effort, or work, has been less frequently considered. The data suggest that !Kung men do not experience subsistence hunting as mere stress, which depresses T levels, for its main effect was an increase in evening values. On the other hand, no interaction of hunt outcome and T levels were observed, in contrast to reports that T reflects success and failure in certain aggressive and dominance interactions. Rather, these data are most congruent with reported effects of moderate prolonged excercise." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 456)

"We conclude that subsistence hunting was associated with significant changes in testosterone patterns in !Kung hunters, due primarily to the maintenance of T concentrations during hunting days that is reflected in elevation of evening values. Diurnal variation in circulating T was hence markedly attenuated. It is also notable that shifts in T level were a consequence, not a primary cause, of the behaviors observed here; hormone changes followed and supported, not preceded, hunting activity. Studies of human behavioral biology might consider the ways in which the dominant context of human evolution, the hunting-gathering niche, may have selected for efficient, appropriate responses to subsistence-related effort." (Worthman, Carol M. & Konner, Melvin, J. (1987) Testosterone levels change with subsistence hunting effort in !Kung San men. Psychoneuroendocrinology 12(6): pp. 456)

"Readers will question why I should attempt reviving such biological heresy long since thought buried. The seasons lie not in Darwin's concept of selective survival but with the neo-Darwinian reliance on two notions which have guided thinking on the subject for about a century. Currently these notions are (a) that the locus of all meaningful phylogenetic changes (in multicellular organisms) occurs only in the germline genes of the gonads, (b) that the genes in the gonads are resistant, with the exception, for example, of ionizing radiation, to most direct influences of the contemporary environment. I do not feel these premises, when coupled to the concept of selective survival, help explain the phenomenon of evolution as we perceive it." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 2)

"The other point, which Mayr's summary illuminates, is that Darwinian selection as perceived in the current paradigm is basicly a negative selection theory. Variant individuals arise in a population and those which by chance are unfit for survival are eliminated by the environment. This ordering process
provides a rational mechanism for guiding evolutionary change to some extent in large populations of individuals, but I feel it is a rather crude conceptual device and unlikely to be the final word on the evolution of multicellular organisms: to enrich evolutionary theory I think it will be necessary to add the complementary, yet opposing, concept of a positive selection force (feedback) also supplied by the environment." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 45)

"If the central dogma is correct, what do we do with Weismann's doctrine? Can it be replaced with something more useful and rational? I am aware of no direct observations which unequivocally prove that genetic information (DNA, RNA) cannot pass, in principle, from differentiated body cells to ova or sperm. There is also no good evidence that it can. I will stick my neck out and predict that future research on the endogenous C-type RNA viruses will very soon supply evidence that C-type RNA viruses can (a) "capture" genes in somatic cells (probably mRNA), (b) cross the tissue barriers partitioning the gametes in the gonads, (c) infect ova or sperm (or travel as extracellular particles in the seminal fluid, infecting the egg around the time of fertilization), and (d) by a process involving reverse trascriptase (RNA-dependent DNA-polymerase), synthesize a DNA copy allowing the integration of these captured somatic genes ("normal" or mutated) back into their appropriate positions in the germline DNA. This step is a crucial feature of the model because it demands a high degree of integration fidelity as to where "new" gentic information is to be positioned to satisfy the four dimensions of future development." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 48)

"The hypothesis states that new adaptive processes begin within the somatic cell populations of individual organisms. Environmental stimuli select and order genetic mutations occurring by chance in somatic cells of a given tissue or organ; if the selection stimulus is intense and chronic, there is an increased probability that these new "adaptive" genes are captured by the appropriate endogenous viral vector and integrated back into their homologous Medelian loci in the germline DNA of that individual. In this way, the organism both adapts to its own environment and anticipates, to some extent, the future environment of its offspring: allowing them to adapt instictively, if this selection process continues. The synthesis of this hypothesis is achieved by fusing two widely acknowledged theories, both of which enjoy considerable experimental support: (1) the clonal selection and somatic mutation theory of antibody production and (2) the proto-virus and provirus hypothesis of the endogenous RNA tumor viruses." (Steele, E. J. (1979) Somatic Selection and Adative Evolution. Chicago: Chicago University Press. pp. 60)