"LYSENKO, VIEWS OF NATURE AND SOCIETY -
REDUCTIONIST BIOLOGY AS A KHRUSCHEVITE REVISIONIST WEAPON"
First published in pamphlet format in Toronto; September 1993. (pp.96-151)
Continuing With: Part 1 - (In Web edition- Section 3 of)
As the role of the nucleus came under intense scrutiny by powerful microscopy, it seemed to reveal partial proof of a profound continuity. The nucleus never arose without a division of a pre-existent nucleus. This observation became the key vehicle for those who proclaimed a "Bearer of Heredity". This became the central basis for those clustered around Weismann : Weismann became a central figure in controversies about the role of acquired inheritance : In rejecting Inheritance of Acquired Characteristics, Weismann proposed a : Mayr proposes that Weissman was basically correct in his supposition that there is a complete separation "of the germ plasm from the phenotype expression in the body." But to bring him up to date with molecular biology, he suggests a modification : However, having disposed of Lamarckism, and the remnants of so called "soft inheritance" in Darwin, Weismann now proposed to erect his own inheritance theory. Ernst Mayr adduces the essential facts of Weismannist theory to be : Of these postulates, Mayr goes on to say : Though actually if it is true that the plastids of plants contain hereditary particles, Mayr errs in rejecting postulate number (3). But nonetheless, even an adherent of Weismann is forced to acknowledge that Weismann did not develop a useful theory. Indeed, his theory was robustly criticised on the grounds it did not fit the observed facts : It is interesting that Mayr acknowledges that Weismann believed ultimately in extreme preformationism : Nonetheless in summing up Weismann, Mayr unreservedly eulogises him : Thus Weismann was the quintessence of a Darwinist.
We have explained that by this is meant that he thought the presence of natural selection was completely adequate for explaining evolution and that he agreed that chance events alone drove variation.

But Weismann's major legacy to his later followers, was the strict "law" that controlled and outlawed, any profligate intercourse between the nucleus and the cytoplasm. This became of course the famous Central Dogma. Like its' iron and metal counterpart, this chastity belt did not fulfil its role in the real world either.

His most commonly cited experiments in the present era, are those that are said to prove that the inheritance of Acquired Characteristics cannot take place.
In these experiments, he cut off the tails of adult mice in several generations and examined the progeny for evidence of tail shortening. When no apparent evidence for a tendency to shortening emerged in the progeny over generations, Weismann concluded that here was no evidence for Inheritance of Acquired Characters.

From early on, a critique of this experiment has been that tail shortening does not contribute to survival values for the mice in this environment. But the experiment is cited repeatedly as proof that Inheritance of Acquired Characters cannot occur.

Even before the end of the century he was to be attacked for his "speculative" division between the soma and the germ plasm. Johanssen led the attack, particularly when Weismann became emboldened and imperial with the coupling to Mendelian theory (See below).

JOHANN (GREGOR) MENDEL AND PARTICULATE BEARERS OF HEREDITY

Mendel (1822-1884) was an Augustinian monk at Brunn (Brno in Czechoslovakia). He presented two papers to the Natural History Society of Brunn on 8 February and 8 March 1865, which were to have a delayed, but ultimately far reaching effects on the genetics. Though his work was published in the Transactions of the Brunn Natural History Society, it remained buried in libraries. An uncut copy of his book was later found in Darwin's library. But the work was basically lost to general view. Until, that is, De Vries, Correns and Von Tschermak, (all of whom were also involved in breeding experiments) re-discovered, and verified his work in the 1890's.

Whereas Kolreuter and others had been breeding and examining hybrids between species for some time, Mendel concentrated upon intra-specific crosses. These time consuming studies, (which for his 1866 papers took eight planting seasons) he analysed tens of thousands, or hundreds of thousands of plants. In so doing, he simplified the problem of analysing changes in heredity. He reasoned that :

Mendel selected peas (pisum sativae) for his experiments because as these were self-fertilising plants, he could keep control of the progeny. Having tested 34 stocks of different varieties that differed in a number of respects, he began his experiments. The following text is aided by the diagram on p. 95, also purloined from Briggs and Walters :

He found similar results for other characters in the pea family. In order to : Thus Mendel had shown definitely that the various characters that make up the phenotype of an organism, were transmitted to the progeny, at least in some cases, by a separation of characters ie.that CHARACTERS SEGREGATED.

This meant that when one character was transmitted or inherited, it did not necessarily get transmitted along with another separate character.

His diagrams and data were also explicitly understood as representing particles, these were so often later to be called, the PARTICULATE BEARERS OF HEREDITY. Since there was segregation, this strongly implied that the characters had to be separable physically.
This led directly to the notion of a physical entity - "elementes" for Mendel, "genes" nowadays.

However one major remaining issue was however, did this explain all features of inheritance?
The orthodox geneticists would maintain that it did.

NON-MENDELIAN INHERITANCE, R.A.FISHER AND GOOD DATA

In general Mendel's laws still have major application to practical genetics, such as counselling parents of some children with congenital defects. However, it is probably an over simplification of a complex situation to apply it to all genetic problems.

From as early as Mendel himself, problems were encountered with the overall theory.
The earliest problem was faced by Mendel himself, who could not replicate his findings in another species, Hieracium. But in general, his theory of segregating factors, or "Elemente" seemed to apply to an impressive list of species, as it was complied by Bateson (See Briggs and Walters, Ibid, p.65).

But two problems in particular, have cropped up with the general extrapolation of Mendelism, to cover all heredity.

The least significant are some allegations of falsification of data.
This is still an open question. Though the undoubted and real significance of whether or not Mendel falsified data - is actually very small.
This is firstly because undoubtedly, Mendelism has a significant explanatory power.
However, his numbers appeared to be "too good", to the doyen of mathematics in genetics, R.A.Fisher. The episode is recounted by Briggs and Walters:

Ernst Mayr points to the fact that as Mendel was trained in statistics and physics, he was a very compulsive experimenter. In which case, there certainly do seem to be oddities, as Mayr explains. Mayr ends by generally exculpating him from scientific mischief. But Mayr does accept that Mendel may have tossed out : Although whether  or not Mendel's theory is discredited, by being open to charges of falisfication is oen matter. On this we stand our ground - that Mendelism does offer some explanatory power and cannot be simply discarded. But, this issue in Mendel's data is potentially a problem in terms of a broad generalisation to a range of other characters and phenotypes. But the real problems of a universal generalisation of Mendelism comes from other sources, such as clear instances of non-Mendelian inheritance.
But there is a different question that arises.
After all, "What is good for the goose is good for the gander too". The point here is that in general a rather "benign" eye is cast on these minor blemishes, upon Mendel who is within the pale. ,
But a rather "harsher judgemental eye" is usually cast upon those outside of the pale for alleged and actual "data re-touching". This includes Lysenko.

A more fundamental problem for the general theory of Mendelism in its domination of genetic theory, which is a problem not reliant on the accusation of falisfication. It remains a serious issue today, especially with the dioscoveries of "maternal inheritance" via the mitochondria (See below)..

Though the phenomena of Mendelian segregation was universally accepted, some of the problems with its universal applicability were noted early on. However, these were not considered very serious. The general view was that they were of no overall relevance to the interpretation of the inviolability of the Central Dogma. These observations were eventually put into a workable framework by the discovery that plastids and chloroplasts contain particulate strands of nucleic acids, that serve as reservoirs of heredity transmission within the cytoplasm.

But of course, this carries serious theoretical consequences for the Weismannists.
Perhaps one of the earliest observations challenging Mendelian segregation as a universal phenomenon was by Carl Correns:

Other instances continued to arise where a convenient Mendelian inheritance pattern was simply not tenable. This includes the inheritance of variegation in Maize uncovered by Rhoades in 1943 : Other examples of similar phenomena tending to an anti-Weismannist interpretation are cited by Briggs and Waters. They comment that: The implications of this are transparently abhorrent to strict Weismannists. However to those not so inflexible they carry some explanatory powers.

As pointed out by Briggs and Walters :

Briggs and Walters link these examples in a continuum from clear non-Mendelian Inheritance (eg. Mirabilis jalapa) to phenotypic variation in plants induced by environmental changes to phenotypic variation as a function of developmental variation and itself able to be triggered by environment. More examples of this are discussed under Lysenko, in Part II.

For now, the accompanying illustration on Page 101 (from Briggs and Walters) serves to underline some plant "misbehaviour" according to strict Mendelism. Thus in Ivy (Hedera helix) there is a marked leaf heterophylly, a variability in leaf shape that reflects a developmental transition in plants called Heteroblastic Development or :

The wild Ivy, shows a lobed leaf only on non-flowering normal flat growing shoots. But on flowering shoots, which are erect, and branch radially, there are only simple leaves. Seedlings are lobed leaved. But portions of either vegetative or flowering shoots can be produced by detaching and growing them, when they can be continued for generations. Of interest, tricks such as repeated cutting, grafting on juvenile stocks, or spraying by gibberelic substance a growth factor-ablates this juvenile state.

FIGURE : Heterophylly in Ivy Hederex.

From, Briggs and Walters; Ibid; p. 101.


That plants may behave in a less than clear cut Mendelian fashion is one thing. But that human disease and inheritance can also be less readily "Mendelian" is more challenging for the Weismannists. But an increasing awareness of so called Non-Mendelian patterns of heredity in man is leading to new approaches to genetic counselling : This development is likely going to force even future modifications of current evolutionary thought.

JOHANNSEN FIGHTS RENEWED WEISMANNISM-MENDELISM

As outlined, Darwin's insistence upon a slow change did not pass unchallenged. The proponents of rapid change were forced to attack Weismann, who became the leading, militant "Neo-Darwinist". The opposition came to be led by Johanssen, De Vries and Bateson :

But in spite of his failure, De Vries inspired others to attempt to do the same, and Morgan set out to do so with Drosophila. Johanssen was scathing about the attempts by Morgan to raise new species on the basis of gene mutations. Johanssen came to believe in "a great central something, as yet not divisible into separate factors" located in the cytoplasm. (Sapp Ibid, p.22). Johanssen disagreed intensely with Weismann, and stated his disagreement within the context of formulating his distinction between the genotype and the phenotype. But this distinction was quite different from the one that Weismann had made. Weismann's chastity belt that was placed as a guard for the genome from the amorous attentions of the cytoplasm, had made a very meaningful distinction.

But for Johanssen, the key question was: What experimental data was there to support this distinction between cytoplasm and nucleus in hereditary matters?
He was profoundly uninterested in any extrapolations from social life and other philosophies. In his view the theories of heredity current then, derived even their language from ideas of transmitting property to the progeny of a family. So he disagreed even with the terms of reference that Weismann used.

Johanssen argued there was no data to support this distinction between the genome and the cytoplasm : And furthermore, he denied that it was possible to have such a distinction between the soma and the germ plasm at all: In fact Johanssen was at bottom, trying to emphasise that Mendelian theory was not the inseparable twin of Weismannist dogma and speculation : Furthermore Johannsen in common with many of the European geneticists fought against a dissection of the genotype for more than descriptive and analytic ends : In arguing so consistently, Johanssen was the precursor of many later biologists who pointed to a dialectical solution of apparent polar opposites in biology. The solutions would not be easy to provide, and in fact are still missing. In confronting complex nature and reality, dogmatic unipolar, or Mechanical Materialist explanations are certainly much simpler to provide.

THE SUICIDE OF PAUL KAMMERER

Kammerer was an Austrian biologist whose work claimed to show hereditary effects of use and disuse, a Lamarckian mechanism. He performed painstaking experiments upon the male midwife toad (Alytes obstetricans). This animal develops nuptial pads in order to grip the female during the mating, but only in the presence of water. The pads are absent in the absence of water.

Paul Kammerer's scientific life was an extraordinary saga that ended in public condemnation by the extremely militant Darwinians, especially Bateson, and subsequently his suicide. The manner in which the academics have treated this tragedy is instructive. Even the more astute and progressive of them have to somehow excuse this shocking episode as a passing abberation, not representative of science. Thus Hull seizes on flimsy evidence of an unhappy love affair, bowdlerizing Koestler's book while citing him!

To add injury to insult, Hull then cites Gould to say that in any case Kammerer was still wrong!

There are two ripostes to Gould here.
One is that at the period under which Kammerer was under attack, his opponents did not point out this reasonable alternative. Thus a logical argument was never presented for Kammerer to attempt to either accept or rebut.

But secondly, Gould's own view allows him to see that the interaction between organism and environment can indeed produce rapid change. Thus he is predisposed to accept that Natural Selection may be responsible for a fast change in Alytes obstetricians. However, the adversaries of Kammerer (old and new) cannot accept that biological change is so rapid. This appears to be the crux upon which the vicious debates occurred.

TOWARDS A CHROMOSOMAL THEORY OF INHERITANCE

Speculation about the role of the nucleus had preceded even its full cytological definition. But before any one could propose a primary role for the nucleus or chromosome, certain information was needed. The individuality of each chromosome, and the existence of the chromosome (though it was no longer visible at this stage) through the cell cycle needed to be established.

This was done by a variety of workers. Already in 1891 Henking had observed that during meiosis in an insect called Pyrrocoris, that half the spermatozoa received 11 chromosomes while the other half received not only these 11 chromosomes but an additional heavy staining body which he called X.

McClung then associated this with the sex chromosome (Mayr Ibid, p.750-751).

Therefore MEIOSIS was detailed and understood in this manner. in fact, another term for meiosis is Reduction Division, and it takes place in the sexual cells of the germ cells in gonads. This "reduces" the chromosome number in the cell by half. Upon fusion, following the sex act, the sex cell or gamete (egg or spermatozoa) fusing with its opposite sex partner gamete cell (ie. either an egg or spermatozoa), will then have the full number of chromosomes.

A corollary of this was that the disappearance of the chromosomes at the beginning of MITOSIS (the division of one cell into two with growth) was only apparent, and not a real disappearance. The structure and visibility of the chromosomes may have changed, but their essential "messages" or chemicals were still there.

It  now became possible for Theodor Boveri in 1891 to state that :

Boveri also, correctly, argued that half of the chromosomes were derived from the father and the other half from the mother.

Montgomery (1901) and Sutton (1902) independently showed that each chromosome was individually recognizable during mitosis and meiosis. They also showed the homology of pairs of chromosomes (that is that the parental and maternal contribution each gave rise to a corresponding partner chromosome, the two together formed a pair).

Furthermore, the demonstration that each chromosome was itself made up of two sister CHROMATID strands, allowed the further understanding that Mendel's "particulate bearers" could be understood as a combination of two genes for each character. That is to say each character was associated with a gene, which came in two doses and types (DOMINANT OR RECESSIVE), called two ALLELES.

Sutton now proposed that :

These early links of characters to a single chromosome inspired the future work of T.H.Morgan whose : But it was probably Barbara McClintlock, working on Zea Mays since 1927; who furnished the proof of the actual physical link between genes and chromosomes.
Her paper, published with Harriet Creighton in 1931, was called : Thus by the beginnings of the 20 th century, some of the planks had been laid for an understanding of the cellular basis of heredity. However, all sorts of factors still did not allow for any adequate sense of how each cell could be in the right place at the right time and doing the right things. This highlighted the problems faced by developmental biologists in particular. Throughout this time, there continued intense debate on the problems posed by the development of organisms. That a single gene concept was inadequate was recognised widely, even by T.H.Morgan, the later unchallenged doyen of the gene theory: This early view of Morgan may relate to his background as an embryologist: THE MORGAN GENE THEORY REDUCES COMPLEXITY

In taking the challenge of a genic explanation further, Morgan resorted to a simple and prolific species, the fruit fly or Drosophila melanogaster. In solving some problems, Morgan followed the Mechanical Materialists, by simplifying and narrowing his vision. This allowed a degree of explanation in an exceedingly complex situation, but fell prey to the reductionist traps of simplification.

T.H.Morgan collaborated in the "Fly Room" at Columbia, New York with A.H.Sturtevant, C.B.Bridges and later H.J.Muller, and proceeded to solve some critical mysteries about the behaviour of the chromosomes.

According to his colleagues, Morgan's single most important contribution was the description of CROSSING OVER. Because each chromosome had two chromatids, and becasue during Meiosis, before the chromosomes "disappeared" from view, they became intertwined in a complex manner, they could and apparently did, leave bits behind as they "crossed". It was easily seen that this allowed some degree of change and variability in the genome. In 1946 Muller, the future Nobel Prize winner, and an interested partner in the field of Soviet Genetics (See Below), commented that this and :

The Fly Room group elaborated the following principles: As discussed above, Morgan had at one stage been a developmental biologist. Until 1926, Morgan would allow for some role for the cytoplasm in cell determination; particularly in the development of organisms. At that stage he recognised that a single gene concept, to explain all phenotypic phenomena was inadequate. Keller is the biographer of Barbara McClintlock who would challenge the supremacy of Morganism in Western genetics, and provide explantions of the enormous "flexibility" of the gene (She coined the term of the so called "jumping genes"). Keller understand the signifcance of Morgan's departure form his previous emphasis on "ontogeny" ie. on development. Now that Morgan's team had used the Fly Room to elaborate their principles, it was expected that all problems of genetics, and indeed of evolution would be solved fairly rapidly. But, despite confident expectations, Morgan's laboratory could not show changes akin to evolution with mutations. This early disappointment was a pointer to a more complex reality. In fact nowadays: Morgan acknowledged that there may be a problem in extrapolating from the bench to true life: But by 1932 Morgan's critics were met by accusations of irrationality and mysticism: The dilemma of the Mechanical Materialists (See parts in previous section on Chance) was reasserting itself.

In a complex world, if you simplify enough, an explanation of some phenomena will surface. It may not be the whole truth. But if you insist on the explanations's inadequacy you will have to develop a more sophisticated explanation. Unfortunately that may take time to develop.

Certainly, by the 1920's most American geneticists had come to view the gene as discrete physical units located on chromosomes. They claimed that genes were the "governing elements" of the cell largely immune from the rest of the cell, yet dictating its activities. T.H.Morgan expressed this view succinctly in 1926:

Current textbooks on molecular genetics suggest an untrammelled development towards today's Orthodoxy. For Ernst Mayr, a key representative of the New Synthesis between Darwinism and Mendelian genetics: But this view was not achieved by quick, or even unanimous approval.
Ernst Mayr himself a founder of the New Synthesis, acknowledges the resistance put up by scientists to the Chromosome Theory of Inheritance.

The resistance itself was a profound uneasiness with structures and processes that seemed to be at odds with notions of plasticity and change. It was precisely for this reason, that the resistance was often most strongly articulated by EMBRYOLOGISTS. Ironically of course, this was the very heritage that Morga himself had warned held the most important problems, and therefore lessons for heredity.
The experiments of Hans Spemann illustrate why embryologists in particular showed such profound resistance.

Spemann had perfected a technique that yielded influential results. He reasoned that he if he could separate the cytoplasm from the nucleus in a developing embryo, he could test the biological function of the two parts:

This seemed to indicate that the nucleus was not able to unalterably change the embryo, in such a manner that the cytoplasm by itself could not also accomplish in the early stages of the embryo.
These various experiments of the merogony school did not unduly perturb the Morganists whose path had been now determined.

It is not surprising that the embryologists were resistant to a narrowly based theory relying only on an unchanging, and fixed master control from the nucleus. After all, the stock in trade of embryologists, was the very antithesis of static structures, as the embryo was ever changing and in an intense flux:

In fact many prominent scientists in the West, have disagreed with the orthodox Morganist school, as will become apparent as the story progresses.

Nonetheless, acting as cheerleaders for Morgan and his Fly Room, was the School of American genetics centred on E.B.Wilson. These scientists consciously wished to establish the chromosomal basis of genetics. This aim faced opposition because it countered the view that it was incorrect to "artificially" split up the organism's functioning:

Naturally the progress of Morgan's group was welcomed by Wilson. But even he, an "invaluable ally" of the chromosomal theory, was forced by 1923 to write: THE NEW SYNTHESIS: The Integration of Evolution and Gene Theories Of Heredity

The field of evoutionary biology had been following the events in herdeitary and genetics clsoely. If anything, they were as much in theoretical disagreements as were the developmental biologists:

Those working to a simplification of the complexities of evolution were attracted to the comparable simplification - with associated huge experimental strides - made by the Morganists. They would join forces later on under the rubric of the "New Synthesis".
In the meantime, the direction of all work seemed to be to preserve the "inviolability" of the genome. This after all, much simplified a conceptual approach as well.

ORTHODOX MOLECULAR GENETICS

Many scientists rapidly developed the further detailed basis of the molecular view of the inheritance of Mendelian characters. But we can only discuss a few key participants. In formulating the new molecular biology, the attacks on Inheritance of Acquired Characters became critical to the new Molecular biology.

This paper, originated in an inspiration at the Faculty Club, where Luria was watching a Slot Machine game. The experiment seemed to forcefully address the issue of whether or not there was a chance variation of organisms, or whether there was non-random, or directed mutations variations, perhaps in response to the environment. We have previously (See p.73) looked at the issue of "chance", or randomness in biology. We will readdress this issue with data provided at a molecular level (See p.132).

But here we will only note that this demonstration by Delbruck and Luria was enormously powerful in reassuring the neo-Darwinists, that the environment could be safely ignored as an inducer of heritable signals.

Of course, the molecular basis of heredity remained a persistent and tantalising question :

"What molecules could transmit heredity? What does a "gene" look like and what is it made of?

Oswald Avery investigated Frederick Griffith's discovery of transformation by dead pneumococcal virulent bacteria of live but avirulent bacteria. This classic experiment became much cited by the proponents of cytoplasmic inheritance (See below), and by those advocating a chemical basis for the gene. Quickly the focus of speculation became the DNA molecule.

The biochemists had become aware of the DNA molecule, and this now concentrated research. By 1944, Avery, Colin McCleod and Maclyn McCarty had found that Griffith's observations was due to DNA. Al Hersey and Martha Chase in 1952 showed that on infection of bacterium by a virus (Bacteriophage) ONLY DNA enters. The focus on DNA seemed odd to many.

Some were sceptical. Delbruck called DNA "a stupid molecule" because it was only composed of 4 bases. How could it specify all the different information required? It was George Gamov, a physician, who saw the problem was a trivial matter of coding. By 1953 Watson and Crick had "cracked the code".

They used the knowledge that the DNA molecule contained equal amounts of pyrmidines and purines, (the four "stupid" bases). They also obtained insight from Rosalind Wilkins who had made X-Ray crystallographic views of the DNA molecule. In 1957 they built on a cardboard model, the molecular structure of DNA. This allowed them to enunciate the "Central Dogma".

Jaques Monod was now able to simply reiterate the old "Weissmann Doctrine", dressed up in the new molecular terms. He therefore now claimed that: By now, it appeared transparently clear, that Weissman was right all along. Indeed it seemed that the nucleus was all that dictated heredity.

EARLY 20 TH CENTURY "UNORTHODOX GENETICS"

Of course this demonstration of the simultaneously intricate, but apparently simple biochemical core of the nucleus; became a major analytic tool. The genetic views expressed above in the Central Dogma certainly did become the dominant ones. Accordingly, most biologists did not stray far from these paths. But molecular orthodoxy itself led to many distinguished non-conformers.

And because of the notable theories they were involved with, we should outline them. T
hese theories in one way or another, tend towards a dialectical realisation that the gene theory as stated was not sufficiently able to grasp reality by itself. The authors would have shuddered to use the term "dialectical", but their views did incorporate many features of dialectics.

In the USA there was Barbara McCLintlock, T.M.Sonneborn, and Ruth Sager. But a great tradition in this area commenced with the German Wissenschaft field, and we explore first the views of Carl Correns, Wettstein,and Michaelis. Their work helped inspire some later USA workers including Sonneborn.

Some members of this German school emigrated to the USA as refugees of the Second World War. Their previous work had brought them fame and scientific credits. But this credit was within an anti-orthodox tradition. As a consequence, their reception in the USA was less than amicable, as seen in the careers of Richard Goldschmidt, and Victor Jollos.

In France, a strong Lamarckian tradition had long held sway, and in the German school, a strong tradition incorporated embryonic development into theories of genetics. The German Wissenschaft tradition, stemming in part from Kant encouraged a strong unifying approach (Sapp, p.59).

As a result, a school with a more holistic view, one that saw a need for both a cytoplasmic view and a nuclear view evolved in Germany. Here the term "Kernmonopol" (Kern = nucleus in German) came to mean that Morgan's points were one sidedly dominant. Though in the USA the Morganists were dominant; in German biology, Jacques Loeb was dominant.

Loeb, in an equal and opposite (and jsut as dogmatic) reaction to Morganism believed that the cytoplasm was the key:

Loeb argued that the cytoplasm was : Loeb's influence was pervasive.
But it was the group around Carl Correns, Richard Goldschmidt, Hans Spemann and Max Hartmann who challenged Morgan-Mendelian genetics fundamentally, by their experimental data.
This group were in the Kaiser-Wilhelm-Insitut fur biologie at Berlin-Dahlem.

Correns had described variegation as a phenomenon of maternal inheritance. When Correns became the Director in Berlin-Dahlem he created a profoundly intense arena for research into cytoplasmic and nuclear interactions. As result of the group's endeavours, a coherent anti-Kernmonopol theory began to take shape.

But this process itself was internally controversial. In this arena, the controversy took the equal and opposite shape from Morganism, and it revolved around:

Following Loeb, Hans Winkler believed that the cytoplasm was composed of self-reproducing genetic entities, dubbing these "plasmagenes". These bore some relation to the  "gemmules" of Charles Darwin. Correns however strongly disagreed with this viewpoint.
Correns preferred to continue to argue for a strictly developmental interaction between nucleus and cytoplasm: This was an essentially dialectic viewpoint. Not that this was a view that was conciously taken however. But nonetheless - it remained alive to the notions of a two-way interaction - the traffic affecting both reciever and giver of genetic information.
ie Both nucleus and cytoplasm have a fundamental interrelationship. Loosely, neither one or the other was by itself adequate. Both were required, and actually the sum of the parts was greater than the individual constituents themselves.

As such this view was in contradistinction to both the Kernmonopolists (Morganists) and the Loebian cytoplasmic forces. This research line of Correns, was taken further by Frits von Wettstein, Correns' assistant. Later as director of the Kaiser-Wilhelm-Institute-fur Biologie, he continued working through World War II. According to Sapp, he was not however a Nazi. His work started from plants, as he realised that the distinctions between germplasm and somatic tissues were much more difficult to make in plants. As Sapp expresses it :

In examining the genetic effects of the cytoplasm, Wettstein followed Correns in stating that these effects could only be examined after crossing genes with very distant genetic constitutions. In many organisms this was not easy, but in some mosses he was able to show profound effects as he had predicted: In order to be absolutely sure of his results, Wettstein set up a highly intricate experimental design. This would test the hypothesis that there were independent cytoplamic effects. But he had to enure that he had: Whilst Wettstein continued his work, Otto Renner had been studying Oenothera in the University of Jena. This had originally provided Johanssen with some puzzles. But : Wettstein now challenged both Morgan and Loeb, for their equla and opposite claims for the heredity hegemony of either the cytoplasm or the nucleus.

He talked of the Idioplasm as the entire genetic capacity of the organism being split up into the genome of the nuclear contribution, and the Plasmon "the structure of the cytoplasm" and the Plastidom representing the plastids in plants. All contributed to the final make up of the progeny:

The baton in German Plasmon work, now passed to Peter Michaelis, after Wettstein's death in 1945. Michaleis, at the Max-Planck-Institut - fur - Zuchtungsforschung in Koln-Vogelsang, carried out further complex backcross experiments. He believed from these that there was a complex unity in the cytoplasm, but he rejected simplistic notions such as the "plasma gene" (so as to "discourage any comparison to the nuclear gene"), or that the cause of cytoplasmic inheritance was solely due to plastids and mitochondria : Michaelis published these views in 1954, and continued to publish right up until 1960. However, in general, all the work of the German school was treated with extreme scepticism and cynicism by the Kernmonopolists in the USA. The embryologists like Ross Harrison, continued to see some need for theories that involved extra-nuclear inheritance. But they were few in number. The careers of Jollos and Goldschmidt are very instructive, in just how antagonistic the dominant majority could be. The dominanat school obstructed the dissemination of the views of dissidents.They controlled after all the journals that scientists need to publish and discuss thier findings. Finally, they obstructed the careers of those with the temrity to question their betters.
Many of these German scientists left Germany for the USA, and their scientific reception left much to be desired.

THE LOCUS OF CYTOPLASMIC GENETICS MOVES:
VICTOR JOLLOS' EXILE IN THE USA

Victor Jollos led some pioneering work in the Kaiser-Wilhelm-Insitut fur biologie, on the effects of environment upon hereditary changes in paramecium, a protozoan unicellular animal. This organism soon became a model for Sonneborn and other American workers, who were heavily influenced by Jollos' work in Germany.

The implications for evolutionary theory were apparent, but Jollos did not wish to make too much of this. He was nonetheless interested in the natural world's potential for changes, in the step-to-step orthogenetic change, that seemd to be required so often - both in development of the individual organism (ontogeny) and evolution of the class to which the single organism belonged (phylogeny). For example in the so called "limb-reduction" in snakes and lizards. Factors in the natural world (as opposed to the laboratory) such as temperature may have really been evolutionarily speaking of interest: About this time Richard Goldschmidt had reported some temperature related experiments: Jollos had by now concluded that indeed Dauermodifikationen could play a serious role in orthogenetic evolution: From the above, it should be no surprise that the pure mathematical modellers, up-holders of the New Synthesis, such as R.Fisher and Sewall Wright were emphatically un-impressed with Jollos' data: It is striking how similarly to Barbara McClintlock (Cited below), Jollos expressed himself on the mathematical school.
Soon however Jollos as a Jew in the middle of a fascist Germany was forced to leave Germany. He accepted a position in the University of Wisconsin in 1934. But surrounded by the school of Kernmonopol - the nuclear domination, things went badly for him from the beginning: The scientific world, especially at these very high academic levels, is very cruel. The image of a "pure science" - uncontaminated by politics is of course, now often recognised as errant nonsense. Enough hoop-la has emerged in recent years, about all the great scientific controversies. But here - an especial political dimension was being touched upon.
            To change or not change?
            "Men of gold for ever - or can men of iron become men of gold?"
The nuclear monopolists followed Plato by refusing that they could. (See Appendix 2).

BARBARA McCLINTLOCK AND THE FLEXIBLE GENOME

And of course, most of these were indeed Men. Women were a rare species in the nuclear monopolised labs. Though later on she became a "genetic" rebel; in her early work McClintlock had proved that there was an actual physical link between genes and chromosomes:

From this discovery McClintlock moved on to work on the Nucleolar Organizer Region of the chromosome. She concluded that changes within the nucleus were highly regulated events. This sense of "cellular and nuclear control", may have aided her later awareness that nuclear events were not purely due to chance. For now however, she was still within the mainstream of genetics. But already she was aware of a greater degree of complexity than the reductionist Mendelians would admit to.

Next she began a life long investigation into the plant zea mays, or maize. She was a meticulous and obssessive recorder. She excelled in accurate painstaking correlations of visible nuclear events with phenotypic and morphological changes. A major step in her development came when she described a novel chromosome. In fact at first, she had purely based such a description upon only deductive reasoning, that was itself forced by an observed - yet otherwise inexplicable phenomenon. The un-described phenomeon, could potentially explain an otherwise unpredictable behaviour in the appearance of leaf striping (variegation). However it was... un-described. But, she then went on to actually visualise it, showing that it did in fact exist. She called it a ring chromosome, because it has fused (ie annealed) ends, which so forms a ring that denies any transcription. Thus its DNA message cannot be expressed. (See Keller p.65-7.)

Exploring further, she showed that the Reannealing (fusing together of broken ends of chromosomes) process was a repair process seen after X-Ray damage, and therefore a predictable, as opposed to a random event :

For our general theme upon links between genome and environment, the significance of these findings lay in the demonstrable flexibility and changeability of the genome : By 1950 she had found evidence of even more complex means of control in the genome : It is not entirely true to say, that in the early years she was completely ignored, despite her discoveries. She did indeed achieve some recognition for her work, being for example elected Vice President of the Genetics Society of America in 1939, a member of the National Academy in 1944, and President of the Genetics Society in 1945. However unquestionably her career suffered due to her manifest heresy, so for example, she experienced difficulty in obtaining a full time position. Her difficulties seemed to be related to her highly individual personality, but also the fact that she was a woman.

Paradoxically her isolation actually helped to keep her insulated from conventional dogma, allowing her creative insight. As her biogrpaher describes it, she "asked questions all her own":

McCLintlock would be in later years, explicit to her biographer about what the New Synthesis meant to her : As Keller points out, the resulting discoveries were heretical in the extreme, beoing "totally at variance with the predominant view of genetics": There was little general appreciation of the relevance of McClintlock's work until much later. It is quite astounding, but in fact, she received only 2 reprint requests for her 1953 paper on transposition! Personal responses to her included phrases such as: Undeterred, teh "old bag", McClintlock went on to describe yet another system of regulation and control, and called it the Suppressor-Mutator (spm) System. This highly dialectical system is simply described by her biographer: All this made the control of genetic systems even more complicated. Certainly the complexity was vastly beyond than envisaged by the Mendelists-single-gene-Morganists.
By now the interaction of the genome with the external chemical environment was becoming demonstrably and palpably a force in genetic discussions: Despite this awareness of the limitations of current theory, the Central Dogma had already become an established philosophical stance.
After this occurred, naturally McClintlock became even more isolated and even more was her work ignored. Apparently unfazed by her relative isolation, McClintlock chafed at limits imposed by established dogma on plasticity of the maize plant: Orthodox geneticists had indeed made suggestions in order to bridge deficiencies, in "patch-work" attempts to incorporate the difficulties, such as those uncovered by McClintlock : Perhaps the two orthodox workers whose work came closest to her were Jacques Monod and Francois Jacob. Interestingly, Jacques Monod, was an ex-Communist Party of France member and hero of the French underground, leaving the CP in 1945. He left it in protests at Lysenkoism. In Keller's words: Mond and Jacob acknowledged the idea of a two-way control, one that included a "feed back", and proposed three fundamental operations, for each of which there shouls be a corresponding gene:
the structural gene, an operator gene adjacent to it, and a regulator gene elsewhere on the chromosome. The 3 together were called an Operon. They wrote: Barbara McClintlock naturally responded positively to this evidence of a move to less in-flexible view of the nuclear genome: McClintlock concluded about their work in a review for American Naturalist that: But rather less generously, and perhaps symptomactic - Jacob and Monod themselves neglected to cite her work in their major paper on regulatory mechanisms of 1961. They excused this later as "an "unhappy oversight", they later called it." (Keller, Ibid. p.10).

Even at this point, however most geneticists thought only about gene transfer mechanisms in bacteria or viruses. Broaching these issues in the complexity of the higher orgnaisms of the eukaryotes, was avoided as McClintlock pointed out:

The neglect of McClintlock's own work had left her quite isolated.
She left Cold Spring feeling "outnumbered", and instead she started field work in Central America on collection of indigenous corn stocks. In this work she contributed to anthropological work on the farming activities of the ancient Americans. (Keller, Ibid. p.181.)

However independent support for McClintlock soon came. In 1963, A.L.Taylor demonstrated that the mu virus (a bacteriophage- or parasite of bacteria) could insert into the bacterial DNA, at many sites:

Shortly after in 1966, Beckwith, Signer and Epstein, acknowledged McClintlock, when they used McCLintlock's term - transposition, in reporting an "F Factor". This was a virus that replicated in bacteria, which could rearrange itself in the DNA. A similar regulation was found in mutants of the bacterium Escherichia Coli.

These mutants themselves inserted some DNA into the sequence of DNA at structural/regulator sites which thereby abolished the function of mutated genes. Upon the precise excision (ie removal of the affected segment), there was a full return of the original (ie non-mutated) behaviour. These were similar responses to the transpositions in maize. Drug resistance phenomena in bacteria usually resulted from phage particles. But a separate mechanism seemed also to be involved where drug resistant genes could move from position to position on a chromosome.

This was sequenced, and bounding the gene on both sides was found repeating sequences - later called a transposon. These repeating sequences were later found to be similar to sequences on the mu insertion element. As Keller remarks:

Peter Stalringer and Heinz Staedler went on to draw an explicit analogy between such phenomenon in bacteria and transposition of maize. They paid her homage by accepting and using her coined term of "Transposable elements". By 1977, this term was accepted widely, thereby offering vindication to McClintlock's vision. As Keller points out, McClintlock was still further ahead in her concepts, invoking jumping genes", and "controlling elements":: But now further work on yeasts and drosophila (on the bithorax complex) have lead to greater acceptance of the notion of "jumping genes". After a lifetime of work in the field, in 1980 McClintlock wrote: This view brought her closer indeed to another Genetic rebel, Richard Goldschmidt (See below).
It is satisfying in this harsh world, that eventually McCLintlock got the recognition that she so richly deserved. McClintlock was asked in 1978 to address the Stadler Symposium on "Mechanisms that rapidly reorganize the genome,". Finally, in 1989, she went on to receive the Nobel Prize for medicine.

But it is by no means fully appreciated even now what the evolutionary significance of theses findings are. Thus Campbell cites a leading evolutionist, Mayr, to show that the evolutionists still have not adequately grappled with a fast changing genome and its implications for evolution :

Why was McClintlock in general so poorly treated by the scientific community?
Her biographer, Keller, adduces several reasons:
the blinkers put on by a pre-existent theory (It can be said that " A Paradigmatic Shift " is required to go beyond the theory), the difficulty of McClintlock's work, the poor delivery by McClintlock - including her own increasingly isolationist tendency ), and the unfamiliarity of zea mays (Keller Ibid p.139-153). Also no doubt, her sex and her unusual personality also played into the reception she received.

But, equally it is clear that a bias against views tending away from the accepted neo-Weismannist view, meant that her and other dissidents were poorly tolerated.

After all, the dissenting views of McClintlock were not unique.
In 1951 Milislav Demerec introduced the Cold Spring Harbour Symposium on " Genes and Mutations " with remarks prompted by three papers by R.Goldschmidt, L.Stadler and B.McClintlock. All three papers focused on mutation. Demerec concluded that:

RICHARD GOLDSCHMIDT, WESTERN UNORTHODOX GENETICS AND SPECIATION

Richard Goldschmidt was a once eminent German biologist, who also became quite an outsider in Western genetics. Like Jollos, he had worked in the Berlin Kaiser-Wilhelm Insitut; and like him had been forced to flee Germany by Nazism. He had begun his attacks on classical orthodoxy in 1933 with his views on speciation. It is true that he was very different from McClintlock in his style of attack on conventional Mendelian genetics.
In short he was very confrontational, and interpreted the available data in a radically different manner from the Morganists.

However, he in fact tried hard to synthesise the nuclear and the cytoplasmic schools, in the process "reconciling embryology and genetics" (Sapp, p. 66). He proposed that the effects of timing of action of the genes was critical.

As so often, it may be said, that Darwin had loosely anticipated this argument. Thus in his "Origin" he had a chapter entitled "Laws of variation", and this contains a Law that Darwin termed;"Correlated variation", by which he means:

Goldschmidt proposed that both the genes and the cytoplasmic interaction could explain much of development: Sapp explains as follows : Though usually interpreted in mere morphological terms, as a purely physical effect, it could be conceived of far more broadly. It might invoke cytoplasmic dominance.
But like the Correns-Wettstein-Michaelis branch of the German school, Goldschmidt rejected the Loeibian view that the cytoplasm determined any hereditary traits by its own action.
He went so far as to say of his own data on the determinants of female sexuality in moths, that he was himself not convinced of the notion of cytoplasmic genes. Instead he explained his data by effects of modification from the cytoplasm. Even this was of course repugnant to the Weismman-Morgan view:: Like Wettstein, he represented a half way house. Yet, even so he was viciously attacked from all sides - by the orthodox, and again by the unorthodox cytoplasm proponents. Lillie, wishing prominence for the cytoplasm, for instance, accused him of a type of Weismann determinism, by invoking genes for all differentiating characters of each stage of development and for: Lillie also charged him with using a circular argument, whereby the cytoplasmic substratum needed to be there at the right time and developmental place, but that its' appearance was determined by the genes.

It seems again in this "piggy in the middle" position, that the more complex Dialectician in biology was "predestined " to be attacked at this period in history.

In fact lately there has been a remarkable resurgence in the thought of Goldschmidt. The concepts of rate limiting genes are now standard in developmental circles.
S.J.Gould uses Goldschmidt's notions in his demonstration of the importance of neonotony, and Dover and Flavell amongst others use Goldschmidt in their views on evolutionary dynamics.

Nowadays, Goldschmidt is most remembered for his views on speciation.
Though he had proclaimed the "gene was dead!", Goldschmidt took the chromosome itself as being the vehicle for generating both change and control of the cytoplasm. Goldschmidt basically argued that since the study of genes dealt wholly with the study of mutations, only inferential deductions had been made about the: "Discrete quantity called the gene."
So he went on, "Why should not evidence of large scale mutagenesis affecting the whole chromosome point to all mutagenic events being exerted on arrangements of chromosomes?"

Goldschmidt felt that recent work (especially McClintlock's) on mutable genes and chromosomal rearrangements vindicated his arguments (in particular). McClintlock's conclusion (in her 1950 paper) that the phenotypic changes she observed resulted not from changes in the actual genes, but from:

Goldschmidt's chromosomes were a fluid and dynamic controller of the interaction between heredity and environment. This allowed the possibility of "sudden" changes in the creation of species. But we have seen how central an idea in the New Synthesis, is the notion of unpredictability, and slowness of change under Natural selection.
Given the extraordinary significance of speciation within biology, it is not surprising that Heterodox biologists should be reluctant to accept the conventional view: For this insight, Goldschmidt's views have recently received more attention. In the theoretical struggle between the theory of neo-Darwinian gradual speciation versus rapid speciation, Goldschmidt has been to some extent "re-discovered".

Goldschmidt's subsequent career, again shows what happened to the majority of the Heretics. Goldschmidt was labelled as being "Obstructionist" by his colleagues. He died, apparently lonely and embittered in 1958:

Despite the similarities between Goldschmidt and McClintlock, she was more reserved about her Heterodoxy, and even had some reservations about Goldschmidt's views: