Final Version May 25, 2003

§7 Mate selection: female choice or Species=Societal consensus?

As described hitherto, one can see that MPR is not a way of mating but a general principle for mating in each species=society. Here we try to verify that the so-called female choice is essentially a species=societal consensus.

In fact it is absurd to suppose: that there were once female giraffes detesting a long neck or peahens disliking splendid tail feathers; but that they disappeared gradually because of their lower fitness. Such a ruminant or such a bird would not have taken part in an incipient species=society of the giraffe or the peacock/hen. In speciation or, precisely, SS partition, the population must have been unanimous concerning its SS symbol.

It is a mating contest in public that decides superior males on the MPR or in sexual selection. Both winner males and loser males are fully aware of their own position in their herd or deme, where inferior males spontaneously retreat from mating activity. For example, male Babirusa pigs in Celebes compete for larger upper tusks. But males having defective upper tusks hide themselves and never appear to the theater of mating contest [Clayton 2001].

An inferior male also respects the result of MPR due to the contest and would never try to challenge for mating, outwitting a superior male. The exclusion of inferior males from mating is admitted by themselves. The usual concept of “female choice” or further “female choosy gene” is unreal because the criterion of selection and the resultant ordering is completely shared and observed by all the mature males and females. Thus sociobiologists will have to create “male choice-suffering genes”. What's more, inferior males in general do not chase females directly but try first to raise their own ranking in the population. They play a role imposed by the law of the SS (MPR) [Mizuhata 2002, 111].

Now it is clear that the so-called “female choice” is an insufficient expression or rather a misconception for a species=societal consensus. Perhaps Darwin, Lorenz, or even Erasmus Darwin must have been aware of this cooperative nature of mate choice, comprising the consensus of males and females (but perhaps R. Fisher and R. Dawkins might have lacked this insight). Yet they were unable to express their correct intuition in correct terminology because they never thought of the concept of species=society and conspecific brain program (or species=societal software).

As it is incorrect to attribute mate selection to “female choice”, its attribution to “female choosy genes” is doubly incorrect: it is a phantom of phantom. As a FCG should correspond to each pattern or trait of each species, FCGs of cichlids must have had to mutate frequently and diverge explosively in advance to speciation while their genoms have remained completely iso-structural. A FCG theorist will have to assert such an enormous fiction for applying their theory to the cichlid problem. And what's more, diverse FCGs seem to have been having similar fitness, where “the difference in fitness” might be unimportant. Anyway multiple fictions are required if one would continue to apply the extant theories to the cichlids.

All these absurdities reveal the fundamental shortcoming of the traditional genic theories. All the complex confusion arises from barren efforts to attribute mate choice, an essentially intentional and cooperative procedure, to a certain unintentional and stochastic mechanism.

§8 Evolution of the cichlids explained by the Neo-Imanishi theory

The observation on the cichlids' evolution gives a marvelous situation to which one can apply the above-mentioned SS theory. The basic methodology here is to treat a speciation as SS partition.

W. J. Dominey and T. Goldschmidt assigned the cichlids' rapid speciation to choosiness of females [Dominey 1984]/[Goldschmidt 1996]. This is half correct and half wrong. It is true that mate choice caused reproductive isolation and hence speciation. However, it may not be correct that they assigned it to female choosiness, or further to the FCG. The true cause for the mate choice is the MPR seeking for SS coherence, which lead to reproductive isolation.

The SS partition corresponding thereto seems to have brought life type partition, and bifurcation in niche preference. As these were all intentional and cooperative processes, they could have got completed in several generations.

As for this rapid SS partition, cichlids seem to have been especially adaptive: the fish seem to have a high visual sensitivity. This character made a prerequisite for quick response to new morphological variants of males. This reasoning was inversely confirmed by the fact that the lake water deterioration and loss of water transparency exert a serious disturbance on all the cichlids' world [Seehausen 1997a], [Mayr 2001].

The cichlids have highly specialized feeding habit and highly adaptive characters thereto [Goldschmidt 1996, chapt.2]. The concerned researchers have interpreted this, hitherto, as a typical case of adaptive evolution as considered by Darwin (and Wallace) [Mayr 2001]. This popular and seemingly “natural” interpretation is, in reality, doubtful: in our theory, it seems to be the same misinterpretation as Darwin's inference for evolution of the giraffe.

Adaptation as the results of evolution does not explain the cause to drive the adaptation, as has been explained for giraffe's evolution. Especially, rapid speciation cannot be interpreted as Darwinian evolution, or unintentional and stochastic “selection”. Giraffe's evolution was driven by their MPR, or more explicitly, their seeking for species=societal coherence. Also the cichlids' evolution must have been driven by their MPR, or more explicitly, their ardent seeking for the species=societal identity and the differentiation from close kin species. This is why their speciation occurred at trivial differences like body patterns and colors, the least concerned with viability or adaptation.

Thus, a SS partition, or a speciation as intentional and cooperative process of the population can occur within several generations, which will accompany the relevant niche specialization or life type differentiation. In this incipient stage, the feeding habit might have changed but the feeding organs, seemingly, had not yet made morphological adaptation. It would have taken far more time to evolve the shapes of mouth, teeth, gill etc. into those more adaptive for eating new foods.

Beware that this aspect of adaptation may not be identical with that considered by Darwin and evolutionists hitherto. There has been found the SS aspect of adaptive evolution: it begins with a SS partition, or intentional and cooperative alteration of life type or mate choice criteria. This precedes and will release the morphological adaptation hitherto considered in the traditional theory of natural selection.

Natural selection might serve to promote this adaptation through unintentional and stochastic discrimination of “the fit and the less fit”. In contrast SS selection, if it occurred, could strongly accelerate adaptation through intentional and cooperative behavior of the population. For example, if females of prawn-eating cichlids prefer to mate with male variants looking adaptive for eating prawns, then this mating behavior (MPR in kind) would surely accelerate adaptation: so to say, this is a “highway” course of adaptation and the Darwinian is a local path. Whether in Darwinian or Neo-Darwinian process, effective adaptation cannot proceed in few generations.

As for the cichlids' species divergence, the unusual speed of speciation itself will reject most usual theories and await a new (unusual?) theory. In contrast SS theory of evolution clarified the motive force toward a frequent and rapid development of reproductive isolation and niche specialization. In conclusion the cichlids' speciation proceeded as followings: mate choice by MPR seeking for a new SS coherence, reproductive isolation, species=societal niche specialization, morphological adaptation through natural and/or SS selection.

One should note that both mate and niche preferences occur as intentional and cooperative phenomena functioning in concert, and not as unintentional and stochastic phenomena by natural selection. The former is essentially a function of the brain [Yoro 2000]. Also remember that members of a SS share the same brain program or the SS software.

§9 Further application of the MPR and the SS partition approach

Analysis of speciation as SS partition can apply, beyond the cichlids' case, to speciation in general because:

1) In any speciation or cladogenesis, the incipient difference between the partitioning populations must be minimal genetically and morphologically.

2) Thus the difference in viability between them must also be small.

3) Therefore natural selection is very ineffective: in fact Darwin expected thousands or tens of thousand generations for speciation.

4) Indispensable for cladogenesis is a mechanism capable to discriminate slight character differences and to “translate” them to mating behaviors (accompanying gene selection).

5) Genic theories of sexual selection are found to be untenable and should be replaced by species=societal theory of evolution.
For example, when the neck of giraffe was still short, the difference (variance) of neck length among males must have been ever small. Yet shorter necked males must have had no less viability because their necks were longer than any female or infant. It is absurd to say that (even inferior) males were more viable or adaptive than females. Thus even adaptive evolution requires some viability-independent motive force. It is not genic but cerebral information processing that can convert slight morphological differences into a decisive discrimination in mating and other ethological behaviors. (The nature and the mechanism of this information processing will be discussed in the next section.)

But for such highly sensitive systems to differentiate peer characters, a viability-independent slight variance cannot lead to disruptive or directive selection. Remember that suppression of visual recognition due to muddy lake water caused a grave disturbance in the world of haplochromine species. Darwinists naively suppose that even a slight character variance could surely cause a corresponding effect, though small yet accumulative, on viability. However, this supposition is dubious. In fact if such a characteristic difference remains “subliminal”, then it would not cause any difference in their ethological behavior and hence in their viability. Natural selection, or unintentional and stochastic selection, is ineffective in principle. Therefore it is hardly applicable not only to the cichlids' speciation but to speciation in more general.

§10 Modes of evolution and the evolution of evolutionary modes

Living things without the brain cannot make selection for themselves. They only suffer selection. Evolutionary “selection” has two modes: an Unintentional and Stochastic (incidental) Mode (is it still selection?) and an Intentional and Cooperative (synergetic) Mode. Here we call the former USM and the latter ICM. (We refrain, hereafter, from using the ambiguous term, “natural selection”.) In USMs, the environment “selects” living things. In ICMs, to the contrary, animals actively (intentionally) select their niche and mating partners, although their options are restricted by the natural environment and their positions in the total biotic community.

　 In the primeval USM, evolutionary “selection” is externally imposed on living things and adaptation is to be caused by the difference in viability (or mortality) among variants or mutants of populations [Kawamiya 2002]. This mode has two aspects: the one (USM1) is selection (preservation) of the fit (or the fitter) and the other (USM2) is selection (elimination) of the unfit. The positive selection USM1 is too tardy for effective scientific proof, as Darwin repeatedly noticed, whereas the negative selection USM2 is well established as had been studied since J. B. S. Haldane and H. J. Muller. @5
@5: The concept of natural “selection” in USM is somewhat self-contradictory because it does not comprise any information processing. Therefore a better term for “selection” in USM would be “screening or sifting” [Mizuhata 2002, 198f]. In Japanese, there have been collateral terms for (evolutionary) selection, i.e. Sentaku (=Choosing) and Touta (=Screening). It is highly advisable to assign USM and ICM to different terms, such as Screening and Selection.　Therefore the traditional “natural selection” should be called “stochastic screening”.
It is noteworthy that the second mode (ICM) had been established only after cerebral animals (which can form species=societies) appeared in the Cambrian Period. The two modes, USMs and ICMs, correspond to the dual layers of the biotic communities: The brainless layer and the cerebral layer. Evolutionists will have to distinguish the two layers because these have a different mode of evolution [Kawamiya 2002]. Cross-layered coevolution is also important as seen in the case of insects and flowering plants. Evolutionists should be aware that evolution after the Cambrian Great Explosion is distinct from evolution before this period [Kawata 2002].

Since the onset of ICMs, it has become difficult to suppose that a pure USM evolution could occur in cerebral animals because the reproductive isolation in speciation occurs essentially in an ICM.

Differentiation of niche preference or mate preference by SS partition (speciation of animal species) belongs to an ICM, in fact it corresponds to revise a version of species=societal software, which comprises the innate releasing mechanism (IRM) and the acquired releasing mechanism (ARM) in animal ethology.

The information processing by the brain in ICMs is considered to be a feedback mechanism of Wiener-Yoshida type [Wiener 1961], [Yoshida 1990a]. It consists of three stages: Cognition (observation/ detection), Evaluation (comparison/ identification), and Direction (decision/ practice). One can easily verify how these stages concern the cichlids' ethological behaviors. For example, species diversity of cichlids occurred in the second (evaluative) stage but their interspecies confusion occurred due to visual disturbance in the first (cognitive) stage. The feedback mechanism comprising the said three stages gives a clear account for the cichlids' behavior in mate choice.

Yoshida asserted to distinguish ICM selection from USM selection. He named the former “Subjective/ Autonomous Selection” against the latter, i.e. usual “natural selection” [Yoshida 1990b]. We highly estimate his original concept created for his “Socio-Information Study”. However, we adopt the term USM or ICM in discussing evolution because these terms are more descriptive.

As seen in the peacock case, one can rather easily identify ICM (like MPR) as such when it functions toward species=society coherent evolution and not toward adaptation. Superficially, however, it is difficult to identify ICM as such when it occurs toward adaptation as seen in the giraffe or the cichlids. Most evolutionists, like Darwin, would take it for an adaptation due to Darwinian natural selection. One should remember Yoshida's remark to distinguish “subjective selection” (ICM) from “natural selection” (USM).

For example all species of cichlids strictly identify their mates by body colors and patterns of males: where the difference lies on the second stage: criteria of evaluation became diverse, meanwhile ways of cognition and decision seem to have been common to all the cichlids.
　 One should note that macroscopic information in the brain cannot be derived from microscopic genetic codes. There is a “Communications Crevasse” between microscopic (molecular) signals and macroscopic (molar) records [Szilard 1929] (just compare electron Volt with micro-Joule. This is an entropy theoretical constraint. In order to get over this crevasse and to convert microscopic signals (DNA or amino acid sequences) into macroscopic information (e.g. brain cell memory), one requires an energy intensive amplifier such as gene analyzers or photo multipliers. Otherwise there is no way to convert molecular signals into macroscopic records. It is strange that most evolutionists are unaware of this thermodynamic constraint and naively lenient to the unproved fiction of “Action Controlling Genes” (ACGs).

Brain cells are sensitive only to macroscopic signals. However, a very round about and indirect interaction may be possible, e.g. genes･･･protein synthesis･･･formation of organs･･･endocrine discharge･･･conditioning of brain activity･･･actual action.

Therefore all the brain memory including its innate part must be transmitted from sources other than genes, where self-organization, interaction with sensory organs and organic chemicals are important in ontogenetic development of the brain [Yoro 2000]. Especially the brain of insects seems heavily dependent on macromolecular chemicals.

§11 Reinvestigating concepts of Species and Species=Society

It is well known that a consistent species concept is hardly attainable as described in the review of J.S.Wilkins: A Taxonomy of Species Definitions...[Wilkins 1997]. The difficulty seems to lie on that the definition has to cover all the species and all their historical transition.

E. O. Wilson defined the biological species as:

A population or series of populations within which free gene flow occurs under natural conditions. What is gene flow? And what are natural conditions? These problems may trigger a chain of problems [Wilson 1988].

As for brain having animals, mating to cause “gene flow” is an intentional and cooperative process described above and a more explicit description of “natural conditions” becomes necessary. Therefore we can define species for cerebral animals as: a population to form a reproductive community sharing the identical Mating Priority Rule (all its modes included).

It should be noted that this species concept has diachronic validity. Human being as a reproductive community itinerated all the vertebral classes: from the Paleozoic vertebrates, proto-fish, proto-amphibians, proto-reptiles, proto-mammals etc., through apes and primates, to Homo erectus and Homo sapiens. Through all these stages, human being must have belonged to the same species. There can be no “origin” for Homo sapiens. Even Pithecanthropus is of the same species as Homo sapiens as far as the former is our monotaxonic ancestor. Even bipedalism cannot sever human being from apes. Identity of human species had continued while they came to live on the ground, descending from trees in forests.
Anyway diachronic identity of Homos can trace back to the proto-vertebrate in the Cambrian Period. In fact the present Homo sapiens must have itinerated through all the classes of vertebrates except the bird: proto-vertebrate, proto-fish, proto-amphibian, proto-reptiles, and proto- mammals in the diachronic anagenesis up to the present human being while retaining their identity as one species. Further consideration leads to the idea that origin of the species must trace back to the origin of life itself. In summary all species exist in a transitional stage, though they seem to belong to each terminal of phylogeny.

As discussed in the former section, evolution of cerebral animals should be treated in ICMs, where species populations form species=society and its speciation or cladogenesis in general occurs as SS partition. Here unit of evolution turns from individuals into species=society [Nature 1985-87]. In parallel to the definition of species, one can resume that: a species=society of cerebral (brain having) animals consists of populations sharing the identical ethological brain program (SS software), which contains Mating Priority Rule as its part [Mizuhata 2002]. And all species=society in concert form the total biotic community [Imanishi 1941]

Species=society has its own history independent of diachronic “gene flow”. The SS software is to be transmitted by intergenerational communication. For example animals learn their MPR through mutual communications in their species-societal life cycle: the MPR is not innate but postnatal (to be acquired ontogenetically). However, as it is a ‘law’ of the SS, it exists prior (prenatal) to each SS individual. If raised being severed from the species-societal life cycle, a giraffe would fail in the practice of the MPR. Therefore, it is not by heredity but by communicative transmission that the MPR is retained diachronically. And this situation is the same for SS software in general.

For example human children raised by wolves acquired wolf's SS software (brain program). Their human genes gave no instruction about how to behave him/herself as a human being [Singh 1942]. After they were retaken into the human society, people tried to teach them language and way of living. However, the children made little progress and failed in acquiring sexuality as human being [Itard 1978]: this means that they could not return to the reproductive community of human being, despite their genes were genuinely human.

In short human beings have no “action controlling genes” (ACGs) by which to live a humane life. However, is it human beings alone that lack ACGs? Do all animals except Homo species have their species ACGs? If so, when and why Homo lost their ACGs? It is time to discard the fiction of ACGs including FCGs (female choosy genes). Evolution theory will no more need such a fictitious hypothesis.

§12 Methodological notes

Evolutionists hitherto seem to have believed in natural selection (in USMs) to be the most essential motive force for evolution and have been trying to reduce an ICM (e.g. mate choice) process down to a USM process. They developed lots of theory on the dubious hypothesis of action controlling genes, whereas what controls animal actions is not genes but the brain program.

In contrast, the MPR is not a hypothesis but a consequence of ethological observations. So ICMs have been widely known since C. Darwin. All we had to do was to identify these concepts as such, setting up correct terminology. These are nothing but Columbus’s egg. This SS approach to evolution will provide an effective methodology for studying evolutions whether adaptive or non-adaptive, rapid or slow, as shown for the cases of the giraffe, the peacock, and the cichlids. The evolution in ICMs is a well-known fact though it has not been identified as such. The MPR theory will replace genic hypotheses concerning mate choice. Especially this new theory can give a clear explanation of speciation process, which had been hardly explained on the concepts of gene mutation.

If ICMs have been equally or more important than USMs, this will reinforce the standpoint of the neutral theory of molecular evolution [Kimura 1988]. Suspicion of panselectionists to the neutral theory seems to have been based not on observed evidences but on their mentality to attribute any evolution to natural selection (USM in kind), where gene mutation controls everything.

That molecular evolution should be neutral (and tardy) is exclusively compatible with the reproductive (genic) continuity indispensable for intergenerational species succession. This genic continuity must have been kept without any leap even at the very instant of speciation. This condition shall be called　“continuity constraint”. Under this constraint one has to elucidate discontinuous phenomena of speciation, where the theory of SS partition will be of great help: discontinuous isolation is completely admissible in ICM evolution　while no discontinuity is admissible in diachronic “gene flow”. Thus the SS approach to evolution and the neutral theory are compatible and complementary. In fact, as seen in the cichlids, a SS partition driven by MPR can explain “congenic speciation”, i.e. congenic reproductive isolation. On the other hand, the neutral theory can explain genic isolation after congenic speciation. If species diversity of the cichlids could continue in LV, without suffering extinction, then they might have genic isolation in future.