重提达尔文的遗传学说——泛生论

长期以来, 达尔文的遗传学说--泛生论(Pangenesis)被认为是错误的。一是泛生论解释的一些现象(如嫁接杂交和获得性遗传)的真实性一直被怀疑, 二是高尔顿的兔子输血实验获得了负面结果, 三是达尔文假定的由细胞释放出来的并可以在体内循环的遗传分子“微芽”(Gemmules)缺乏实验证据。但近年科学文献中积累了许多支持获得性遗传和嫁接杂交的实验证据。循环核酸的发现则说明生物体内确实存在可以在细胞间移动的遗传物质。文章简要介绍达尔文的泛生论及其被湮灭的原因, 并结合自己的工作, 介绍支持泛生论的新证据及我们对泛生论的再认识。重新认识达尔文的泛生论, 对遗传学、医学和进化生物学等领域都有重要意义。     

A new perspective on Darwin's Pangenesis

In 1868 Charles Darwin proposed Pangenesis, a developmental theory of heredity. He suggested that all cells in an organism are capable of shedding minute particles he called gemmules, which are able to circulate throughout the body and finally congregate in the gonads. These particles are then transmitted to the next generation and are responsible for the transmission of characteristics from parent to offspring. If any cells of the parent undergo changes as a result of environmental change, they will consequently transmit modified gemmules to their offspring. Soon after Darwin's pangenetic theory was published, Francis Galton designed a series of blood transfusion experiments on differently pigmented rabbits to test its validity. He found no evidence in support of the existence of Darwin's gemmules and the concept of Pangenesis was largely abandoned. In this article, recent reports of successful induction of heritable changes by blood transfusion are reviewed. Detection of circulating nucleic acids and prions in plant sap and animal blood is considered as fresh evidence for the existence of gemmules. It is now apparent that a considerable revision of views on Darwin's Pangenesis must occur before a new comprehensive genetic theory can be achieved.     

Darwin’s contributions to genetics

Darwin’s contributions to evolutionary biology are well known, but his contributions to genetics are much less known. His main contribution was the collection of a tremendous amount of genetic data, and an attempt to provide a theoretical framework for its interpretation. Darwin clearly described almost all genetic phenomena of fundamental importance, such as prepotency (Mendelian inheritance), bud variation (mutation), heterosis, reversion (atavism), graft hybridization (Michurinian inheritance), sex-limited inheritance, the direct action of the male element on the female (xenia and telegony), the effect of use and disuse, the inheritance of acquired characters (Lamarckian inheritance), and many other observations pertaining to variation, heredity and development. To explain all these observations, Darwin formulated a developmental theory of heredity — Pangenesis — which not only greatly influenced many subsequent theories, but also is supported by recent evidence.     

The History and Reception of Charles Darwin’s Hypothesis of Pangenesis

This paper explores Charles Darwin’s hypothesis of pangenesis through a popular and professional reception history. First published in The Variation of Animals and Plants under Domestication (1868), pangenesis stated that inheritance can be explained by sub-cellular “gemmules” which aggregated in the sexual organs during intercourse. Pangenesis thereby accounted for the seemingly arbitrary absence and presence of traits in offspring while also clarifying some botanical and invertebrates’ limb regeneration abilities. I argue that critics largely interpreted Variation as an extension of On the Origin of Species by Means of Natural Selection (1859), while pangenesis was an extension of natural selection. Contrary to claims that pangenesis was divorced from natural selection by its reliance on the inheritance of acquired characters, pangenesis’s mid nineteenth-century reception suggests that Darwin’s hypothesis responded directly to selection’s critics. Using Variation’s several editions, periodical reviews, and personal correspondence I assess pangenesis popularly, professionally, and biographically to better understand Variation’s impact on 1860s and 70s British evolutionism and inheritance.     

Darwin and Mendel: who was the pioneer of genetics?

Although Mendel is now widely recognized as the founder of genetics, historical studies have shown that he did not in fact propose the modern concept of paired characters linked to genes, nor did he formulate the two "Mendelian laws" in the form now given. Furthermore, Mendel was accused of falsifying his data, and Mendelism has been met with scepticism because of its failure to provide scientific explanation for evolution, to furnish a basis for the process of genetic assimilation and to explain the inheritance of acquired characters, graft hybridization and many other facts. Darwin was the first to clearly describe almost all genetical phenomena of fundamental importance, and was the first to present a developmental theory of heredity--Pangenesis, which not only greatly influenced many subsequent theories of inheritance, particularly those of de Vries, Galton, Brooks and Weismann, but also tied all aspects of variation, heredity and development together, provided a mechanism for most of the observable facts, and is supported by increasing evidence. It has also been indicated that Darwin's influence on Mendel, primarily from The Origin, is evident. The word "gene" was derived from "pangen", itself a derivative of "Pangenesis" which Darwin had coined. It seems that Darwin should have been regarded as the pioneer, if not of transmissional genetics, of developmental genetics and molecular genetics.     

Lamarckism and epigenetic inheritance: a clarification

Since the 1990s, the terms “Lamarckism” and “Lamarckian” have seen a significant resurgence in biological publications. The discovery of new molecular mechanisms (DNA methylation, histone modifications, RNA interference, etc.) have been interpreted as evidence supporting the reality and efficiency of the inheritance of acquired characters, and thus the revival of Lamarckism. The present paper aims at giving a critical evaluation of such interpretations. I argue that two types of arguments allow to draw a clear distinction between the genuine Lamarckian concept of inheritance of acquired characters and transgenerational epigenetic inheritance. The first concerns the explanandum of the processes under consideration: molecular mechanisms of transgenerational epigenetic inheritance are understood as evolved products of natural selection. This means that the kind of inheritance of acquired characters they might be responsible for is an obligatory emergent feature of evolution, whereas traditional Lamarckisms conceived the inheritance of acquired characters as a property inherent in living matter itself. The second argument concerns the explanans of the inheritance of acquired characters: in light of current knowledge, epigenetic mechanisms are not able to drive adaptive evolution by themselves. Emergent Lamarckian phenomena would be possible if and only if individual epigenetic variation allowed the inheritance of acquired characters to be a factor of unlimited change. This implies specific requirements for epigenetic variation, which I explicitly define and expand upon. I then show that given current knowledge, these requirements are not empirically grounded.     

The Darwin of pangenesis

The Darwin of pangenesis is very much another Darwin. Pangenesis is Darwin's comprehensive theory of generation, his theory about all sexual and asexual modes of reproduction and growth. He never explicitly integrated pangenesis with his theory of natural selection. He first formulated pangenesis in the 1840s and integrated it with the physiology, including the cytology, of that era. It was, therefore, not consilient with the newer cytology of the 1860s when he published it in 1868. By reflecting on the role of pangenesis in Darwin's life and work, we can learn to take a wider view of his most general theorising about animal and plant life.     

Synergy Between Low Temperature and Dessication in Breaking Gemmule Diapause of Eunapius fragilis (Leidy)

Summary

Gemmules of Eunapius fragilis collected during the fall and kept at 20° C for up to 6 months did not germinate. Freshly collected gemmules, which were dried at 20° C for 7 days and then rehydrated, also exhibited a very low capacity for germination. However, gemmules, stored at 20° C for several months and then dried, showed a much higher level of germination (but usually no more than 50%) after they were returned to pond water. Gemmules, stored at 4–5° C for 4 to 6 weeks, exhibited at most very little germination when they were tested at 20° C. On the other hand, gemmules, which were chilled at 4–5° C for 4 to 6 weeks and then dried for 7 days, underwent rapid and nearly complete germination upon rehydration. These results provide clear evidence for a synergistic effect between low temperature and desiccation in breaking gemmule diapause. It is suggested that in temporary habitats where E. fragilis often survives the dry summer as gemmules, drying may be the primary agent releasing the gemmules from diapause so that they germinate in the fall upon the return of water. A brief exposure of the gemmules to low temperatures before and/or during the dry period may enhance the effect of desiccation.     

Gemmules of sponges from a warm lake

1. Two types of gemmules were found, each in a different species of sponge, from the warm monomictic Lake Kinneret: (i) clustered gemmules, sharing the pneumatic layer of the gemmular capsule and resembling gemmules of Eunapius ; (ii) gemmules that develop non-synchronously, containing amphidisc spicules within the gemmular capsule and resembling those of Ephydatia . Algal cells were not detected within either type of gemmule although they exist in the developed sponges.
2. Sponges began producing gemmules in the lake with the onset of lake drawdown and ceased when lake level was minimal. The gemmules hatched when the lake level began to rise.
3. Under experimental conditions gemmules hatched between 13 and 35 °C. Germination was optimal at 20–25 °C. Chilling of young gemmules prior to incubation at 25 °C improved germination rate.
4. The percentage of germinating dry gemmules diminished 4–6 months after their collection from the lake. None germinated after 10 months. Submerged gemmules maintained high viability with ageing (up to 100% germination 18 months after collection). Desiccation influenced gemmule viability over time, by both decreasing the percentage of germinating gemmules and increasing the lag time before onset of germination.
5. Gemmules kept in the dark germinated significantly less than those illuminated for 12 h day     

Life-cycles of lotic populations of Spongilla lacustris and Eunapius fragilis (Porifera: Spongillidae)

SUMMARY. The life-cycles of green and white morphs of the freshwater sponge Spongilla lacustris were examined in the light of past evidence that zoochlorellae may augment their sponge host's nutrition. Field collections from a lotic population of S. lacustris were supplemented by laboratory experiments on gemmule hatching and gemmule size. Both white and green S. lacustris produced sperm for a 6-week period in 1976 starting in the middle of May. Out of thirty white and thirty green sponges examined during this period, twenty white and ten green sponges contained sperm. Sperm production in both morphs was limited primarily to the basal 3.18mm of sponge tissue, and the density of sperm packets in the two morphs was the same. Out of 180 white and green sponges examined in 1976, only four eggs, no embryos, and no larvae were observed. White sponges gemmulated a week or two earlier, and produced smaller gemmules which were more uniform in size than those of green sponges. White and green gemmules hatched synchronously in the spring. In 1977 one female and numerous male specimens of S. lacustris , and numerous females but no males of another sponge, Eunapius fragilis , were found. The life-cycles are discussed in the light of other recent studies on freshwater sponges.     

Gemmules of sponges from a warm lake

1. Two types of gemmules were found, each in a different species of sponge, from the warm monomictic Lake Kinneret: (i) clustered gemmules, sharing the pneumatic layer of the gemmular capsule and resembling gemmules of Eunapius ; (ii) gemmules that develop non-synchronously, containing amphidisc spicules within the gemmular capsule and resembling those of Ephydatia . Algal cells were not detected within either type of gemmule although they exist in the developed sponges.
2. Sponges began producing gemmules in the lake with the onset of lake drawdown and ceased when lake level was minimal. The gemmules hatched when the lake level began to rise.
3. Under experimental conditions gemmules hatched between 13 and 35 °C. Germination was optimal at 20–25 °C. Chilling of young gemmules prior to incubation at 25 °C improved germination rate.
4. The percentage of germinating dry gemmules diminished 4–6 months after their collection from the lake. None germinated after 10 months. Submerged gemmules maintained high viability with ageing (up to 100% germination 18 months after collection). Desiccation influenced gemmule viability over time, by both decreasing the percentage of germinating gemmules and increasing the lag time before onset of germination.
5. Gemmules kept in the dark germinated significantly less than those illuminated for 12 h day     

Is cultural evolution Lamarckian?

The article addresses the question whether culture evolves in a Lamarckian manner. I highlight three central aspects of a Lamarckian concept of evolution: the inheritance of acquired characteristics, the transformational pattern of evolution, and the concept of directed changes. A clear exposition of these aspects shows that a system can be a Darwinian variational system instead of a Lamarckian transformational one, even if it is based on inheritance of acquired characteristics and/or on Lamarckian directed changes. On this basis, I apply the three aspects to culture. Taking for granted that culture is a variational system, based on selection processes, I discuss in detail the senses in which cultural inheritance can be said to be Lamarckian and in which sense problem solving, a major factor in cultural change, leads to directed variation.     

Overheated Rats,Race, and the Double Gland: Paul Kammerer,Endocrinology and the Problem of Somatic Induction

In 1920, Eugen Steinach and Paul Kammerer reported experiments showing that exposure to high temperatures altered the structure of the gonad and produced hyper-sexuality in “heat rats,” presumably as a result of the increased production of sex hormones. Using Steinach’s evidence that the gonad is a double gland with distinct sexual and generative functions, they used their findings to explain “racial” differences in the sexuality of indigenous tropical peoples and Europeans. The authors also reported that heat induced anatomical changes in the interstitial cells of the gonad were inherited by the heat rats’ descendants. Kammerer used this finding to link endocrinology to his long-standing interest in the inheritance of acquired characteristics. The heat rats supported his hypothesis that the interstitial cells of the double gland were the mechanism of somatic induction in the inheritance of acquired characteristics. The Steinach–Kammerer collaboration, Kammerer’s use of Steinach’s “puberty gland” to explain somatic induction, and his endocrine analysis of symbiosis reveal Paul Kammerer’s late career attempt to integrate endocrinology and genetics with the political ideals of Austrian socialism. With them he developed a bioethics that challenged the growing reliance on race in eugenics and instead promoted cooperation over competition in evolution. I relate his attempt to the controversies surrounding the interstitial cells, to the status of extra-nuclear theories of heredity, and to Kammerer’s commitment to Austromarxist social reforms during the interwar period. I am very grateful for the help of several archivists, including Valerie-Ann Lutz, Roy Goodman and Robert Cox at the American Philosophical Society Library, Arlene Shaner at the New York Academy of Medicine, Shawn Wilson at the Kinsey Institute Library, the staff at the Archives of the University of Vienna, and Yukiko Sakabe at the Austrian Academy of Sciences. Thanks are also due to Andreas Lixl, who gave very helpful advice on German language and to many colleagues, including three anonymous reviewers, Paul Silvia, Alyce Miller, and the late Gilbert Gottlieb, who provided valuable comments as readers of earlier drafts. My discussions with Veronika Hofer have been especially rewarding. The research was funded by National Science Foundation grant #0240151.     

Pione lampa,a bioeroding sponge in a worm reef

A bioeroding sponge was found in a sabellariid worm reef in Florida, USA. The sponge was investigated with standard techniques for species identification (tissue digestion, microscopy techniques, examination of spicules) and was identified as the clionid Pione lampa (De Laubenfels, 1950). This species is extremely similar to Pione vastifica (Hancock, 1849). Morphological and ecological similarities and possible dissimilarities are discussed. Arguments for conspecifity were found to be weaker than evidence of P. lampa and P. vastifica being separate species. The sampled sponge contained asexual reproductive elements: superficial buds and internal gemmules. Buds were very delicate, consisting mainly of a radially-oriented mesh of collageneous threads. They further comprised abundant granular material and acanthose microrhabds. Numerous choanosomal gemmules were found in the sponge body attached to erosion chamber walls. They were subspherical to lenticular and had an aspicular, unstructured, smooth and rigid coat. It enclosed dense cell material and various spicule types. Whereas buds are interpreted to function as dispersal elements, gemmules will primarily ensure survival under adverse conditions such as smothering, exposure to air and high temperatures. Gemmules from this site might occasionally be freed and scattered, since the Florida reef can suffer heavy damage during periods of increased wave activity such as that created during hurricane season. Bioerosion activity of the sponge increases the chance to free gemmules, as the sponge not only etches into calcareous particles cemented into the matrix produced by the worms, but also into the matrix itself. This ability enables the sponge to utilise the reef as substrate. Within the Clionidae, Cliona annulifera and three species of the genus Pione are the only species known to produce typical gemmules. Possible reasons are: (1) reproduction of bioeroding sponges is understudied and asexual bodies may have been overlooked in other species, and (2) clionid gemmules are an adaptation to survive life in risky environments. Sponges of the genus Pioneare comparatively successful in environments in which they are close to their physical limits or in potentially unstable or mobile substrates.     

Germline epigenetic inheritance: Challenges and opportunities for linking human paternal experience with offspring biology and health

Recently, novel experimental approaches and molecular techniques have demonstrated that a male's experiences can be transmitted through his germline via epigenetic processes. These findings suggest that paternal exposures influence phenotypic variation in unexposed progeny–a proposal that runs counter to canonical ideas about inheritance developed during the 20th century. Nevertheless, support for paternal germline epigenetic inheritance (GEI) in nonhuman mammals continues to grow and the mechanisms underlying this phenomenon are becoming clearer. To what extent do similar processes operate in humans, and if so, what are their implications for understanding human phenotypic variation, health, and evolution? Here, we review evidence for GEI in human and nonhuman mammals and evaluate these findings in relation to historical conceptions of heredity. Drawing on epidemiological data, reproductive biology, and molecular embryology, we outline developments and opportunities for the study of GEI in human populations, emphasizing the challenges that researchers in this area still face.     

Survival of the gemmules of Anheteromeyenia ryderi (Potts) following aerial exposure during winter in New England

Gemmules of Anheteromeyenia ryderi survived 24 h exposure to air temperatures as low as –20 °C under laboratory conditions. Drying the gemmules of A. ryderi at 5 °C under laboratory conditions resulted in a reduced viability and a slower germination rate following rehydration compared with undried control gemmules. Only 25% of the gemmules germinated after drying for one month. Up to 25% of the gemmules of A. ryderi that were tested survived aerial exposure from early November to early April when a pond in Connecticut inhabited by this sponge was drained. During this period air temperatures dropped to as low as –16 °C. Continued aerial exposure of the gemmules during the summer resulted in nearly complete gemmule mortality.     

Darwin and sexual selection: One hundred years of misunderstanding

Darwin's book on the Descent of Man and Selection in Relation to Sex (1871) is often viewed as the continuation of TheOrigin of Species published 12 years earlier (1859), both because of the implicit parallelism between natural selection and sexual selection, and because Darwin himself presents the book as developing a subject (man) which he intentionally omitted in the Origin. But the Descent can also be viewed as the continuation of his book on Variation published three years earlier (1868). Firstly because Darwin's hypothesis of pangenesis links the selection process to the origin of variation through use and disuse, an idea underlying his speculations on the origin of moral sense in humans. Second because like the action of the horticulturist on his domestic crops, sexual selection exerted by one sex on the other sex can develop fancy traits that are not easily accounted for by their utility to the selected organism itself, such as artistic taste, pride, courage, and the morphological differences between human populations. These traits are difficult to reconcile with pangenesis. They add up to other contradictions of the book possibly resulting from Darwin's erroneous inference about the mechanism of inheritance, like those on the determination of sex-ratio, or the confusion between individual adaptation and the advantage to the species. These inconsistencies inaugurate a weakening of the Darwinian message, which will last 50 years after his death. They contributed to the neglect of sexual selection for a century. Darwin however maintained a logical distinction between evolutionary mechanisms and hereditary mechanisms, and an epistemological distinction between evolutionary theory and Pangenesis hypothesis. In the modern context of Mendelian genetics, Darwin's sexual selection retrospectively appears as luminous an idea in its pure principle as natural selection, even though the mechanisms governing the evolution of sexual choice in animals remain largely unresolved.     

Freud’s Lamarckism’ and the Politics of Racial Science

This article re-contextualizes Sigmund Freud’s interest in the idea of the inheritance of acquired characteristics in terms of the socio-political connotations of Lamarckism and Darwinism in the 1930s and 1950s. Many scholars have speculated as to why Freud continued to insist on a supposedly outmoded theory of evolution in the 1930s even as he was aware that it was no longer tenable. While Freud’s initial interest in the inheritance of phylogenetic memory was not necessarily politically motivated, his refusal to abandon this theory in the 1930s must be understood in terms of wider debates, especially regarding the position of the Jewish people in Germany and Austria. Freud became uneasy about the inheritance of memory not because it was scientifically disproven, but because it had become politically charged and suspiciously regarded by the Nazis as Bolshevik and Jewish. Where Freud seemed to use the idea of inherited memory as a way of universalizing his theory beyond the individual cultural milieu of his mostly Jewish patients, such a notion of universal science itself became politically charged and identified as particularly Jewish. The vexed and speculative interpretations of Freud’s Lamarckism are situated as part of a larger post-War cultural reaction against Communism on the one hand (particularly in the 1950s when Lamarckism was associated with the failures of Lysenko), and on the other hand, against any scientific concepts of race in the wake of World War II.     

Microbiome‐Germline Interactions and Their Transgenerational Implications

It is becoming increasingly clear that most, if not all, animals and plants are associated with a diverse array of resident gut microbiota. This symbiosis is regulated by host‐microbiome interactions which influence the development, homeostasis, adaptation and evolution of the host. Recent evidence indicated that these interactions can also affect the host germline and have a potential of supporting transgenerational effects, including inheritance of acquired characteristics. Taken together, the influence of gut bacteria on the host soma and germline could potentially give rise to emergent phenotypes, which may be partially inherited by three distinguishable modes of transgenerational influence of gut bacteria: 1) “soma‐to‐soma” 2) “soma‐to‐germline” and 3) “soma‐germline‐soma”. Here, we discuss these possibilities in light of evidence supporting bacterial‐mediated modes of transgenerational inheritance.