Recapitulating Social Selection sensu Roughgarden through a Mayrian teleonomy of somatic programming

Philosophy and Biology Social selection and sociality in turtles: playing with teleonomy
Here I design a theoretical model of vestibular-sound signaling for turtles in which the empirical hypothesis about the role of social pleasure in enabling teamwork under social selection can be tested. It provides a morphological basis for vestibular signaling as a stable adaptive trait within evolutionary game theory and shows that the notion of strategy vs preference is specious. Thus one becomes able to answer the question (Roughgarden 2012) if individual turtles attain the NBS by jointly maximizing the Nash product or individually maximizing the separate payoffs.

Social selection sensu Roughgarden offers the evolutionary biologist the option of constructing evolutionary theory (forms of social evolution) from the bottom up (2012). This is a new theoretically enabled two tier construction process in which behavior creates dynamical structures in which evolution operates (2006, 2011,2012). Here I suggest applying Mayrs 1961 notion of a behavior computer of an individual to models of social selection in turtles.

This application of ontogenetic irreversibilities that fashion social infrastructure through teleonomic goal-directedness enables empirical decisions on what kinds of games: (competitive-cooperative), repetitive back and forth negotiations, individualistic threats, (NBS

vs NCE) etc. are responsible for geographic differences in relevant trait acquisition. Interestingly, it appears that social selection is responsible for selections of particular mathematically constant adaptive features that can be mechanically understood as a behaviorally motivated goal-directed process that explains some recapitulations without necessarily needing heterchrony to understand the particular relationships between ontogeny and phylogeny between the behavioral tier and evolutionary tier. A dispute between Ayala and Mayr is devolved to the factuality of social selection in populations of constant vs. growing sizes. Programmatic materiality is realized in the proximate reward sensation that has a mathematically compassed spatio-temporality in which the informational metaphor is sedimented by the 7, 1-Dimensional frieze patterns.

by Mark Witton Despite increased exposure (2011,2012)

the notion of social selection sensu Roughgarden, continues to be opposed rather less on the merits than it could be (of a different empirical set of claims than sexual selection). Even within the broader context of sexual selection as a subcategory of social selection sensu West-Eberhard , social evolution through offspring infrastructure sociality is at best considered as fringe (Rubenstein) and at worst rejected outright ( Tobias ,Montgomerie) as simply not related.

This a priori restriction comes in part from a-failure-to-recognize how social infrastructure selection can provide new theoretical probes to unconfound certain physiological mechanisms from particular evolutionary histories. Maynard Smith (J. theor. Biol. (1974) 47, 209 -221 The Theory of Games and the Evolution of Animal Conflicts extending the idea in Smith and Price (1973)which did not consider memory)) only introduced intentionally competitive game theory into the evolutionary biology of ritualized behavior. Lewontins larger provision of organism environment interaction was taken to contrast - not simply within a simple difference of a focus on natural selection (Potochnick Modeling social and evolutionary games 2012) - but was a move to represent larger aggregated (dimensionally greater) ontological entities subject to the same natural selection object and was not pursued then.

and thusly, the possibility of social cooperation within the difference of the apparently competitive difference between a behavioral tournament and a display was modeled strategically otherwise preferring competition as the concept behind the activity or as an implied result of multi-level selection (Gould (after the fact)). In the event it was not stable. Potochnick has filtered this difference in emphasis into claims about the phenotypic instantiation of the equilibrium suggested behaviors. She asserts that game strategies may not be evolved phenotypes in themselves. Instead she sees them as, game consequences of socially supported nonrationality. If{ in some details} however the game is representable inside extant physiological structures functionality (whether or not the notion of limited war applies to snakes that wrestle without fangs or not) then the resultant biomechanics is operative from within the phenotype not globally outside it! Electric fish may use air bladders in their inner ears for other-regard, salamanders may use pheromones as side-payments and head bobbing with turtle sound may bind partially perfect teams into co-incident self- interested coalitions . It is not then derivable from the environment persay but the organismenvironment co-construction (in this instances with memories residing in behavior computer somatic programs). So while it is indeed -- true, that, populations and not individuals have the capacity to evolve, population thinking can be gainsaid by behavioral dynamics under homogenous kinematics of individuals performing different actions. And thus changes in the composition of the/a population that has differences in strategy can be induced empirically from observations of differences of the same individuals themselves performing different actions in some cases. Social selection provides a framework for numerous scenarios in behavioral ecology that remain to be explored. I show how the use of somatic programs as repositories/memories of former threat points abandoned, reclaims the game strategies as particular evolved phenotypes recapitulatable in phylogeny and is a vantage point that can be gainsaid reciprocally between two tiers of social selection sensu Roughgarden. Cooperation rather than competition can be seen to be the drawing force for what appears otherwise as

a moderated conflict. Parrallel driving force minimizations of somatic payoff memories constrain the unconditional use of preferences under Nash Bargaining which display the threat point on the other side of this boundary. Social selection is a purely individual level evolutionary biology constructable. Along with Smith and Price it does not consider snake wrestling or deer antler locking as resulting from some kind of group selection (for the good of the species) but it also does not see behavior conventionally from the human rational standpoint. It attempts to consider the emotional determinates of the behavior rather than the necessarily stable conventionalism of strategies where no mutant can invade. It does strive to find universals for uniform population behaviors. Thus so --- It does not need to introduce higher levels of selection within its praxis from observation to experiment to observation once again (etc.)

Here I explore explaining explicitly a rather imaginary scenario for social evolution in pterosaurs that contains sexual selection and social selection claims and show how there continues to be no reason to reject social selection before data to support or reject it are more fully collected. Details on how to do this with turtles is provided. Social selection praxis is shown to be able to construct evolutionary theory fully in competition with sexual selection and the value of game theory in biology is shown to moderate more extreme claims in the application back into the social sciences themselves In social selection the life-time fitnesses are summed and the deme fitness may depend thus on the ordinality of the invidiuals as they are summed especially if these fitnesses are not commutative with respect to the order of the summation. If this ordinality can be decided as to how it affects the cardinality of the measureable quantities then the social evolution of particular behaviors may affect the gene frequency of any given population as a whole. A restricted range of behaviors would thus exist for a particular set of populations. Thus while it is indeed the case that in reality it is the population that changes , the dynamics of the behavior can be used to coordinate the difference in gene frequencies given an actual determination of the ordertype linking the cardinal and ordinal enumerations of the same effect. This will depend on the probabilities transitioning between individual and team play (Roughgarden et al 2006, August Guang 2012 (Switiching between Cooperation and Competition in Social Selection). This transitioning depends on how the Nash axioms are materially manifested, instantiated and programmed. The quaternionic representation of vestibular functionality provides scale independence to affine transforms (Nash axiom 1). The neurological difference of computation with dual or single quaternions enables the symmetry condition to exist or not (Nash axiom 2). A to be

defined configuration of quaternionic space relative to changing 1-d Symmetries guarentees (will guraentee ) AXIOM 4 independence of irrelevant alternatives)(if a solution has an outcome then a subset has an outcome solvable)). In this generalization of Nash barraging to behavior (animals are not assumed to bargain as humans do (Roughgarden 2012)), behavior computers as somatic programs may, if they serve as memories of threat points abandoned (can possibly )encode the orderytpes of this reflected prior preference and determine future strategy domains provided the physiological manifestations are decoupled kinematically from the evolutionary contingencies that gave rise to the particular individual fitnesses themselves (a broader prior range). Thus there is no theoretical contest between conceiving of game theory in biology during the attainment of equilibrium as preferences or stratigies . That will depend. Rather there are various ways that the Nash Baragining axioms can be organized by different monophlyetic lineages. There may be some where the physiology itself circumscribes Nash Axiom 3 (pareto-optimality). Here this axioms substantial existence is a variable in the behavior computers goal-directedness. The fruitfulness of social selection is found in its providing a plurality of game theory phenotypes involved in optimizing life history strategies that underlies social infrastructure selection. Confusions about the theoretical and empirical boundaries of the process of social evolution are cleared up to some extent through the use of somatic program teleonomy to bridge between the two tiers in some cases (where global teleomatics individualized can be rendered impotent) Ernst Mayr developed the difference between proximate and ultimate causation in biology (from 1961 on ) but some have challenged this distinction which relies for its truth on Pittendridgehs need to unconfound the immediate physiology from the evolutionary history. Social selection sensu Roughgarden with its implicit construction via two tiers (behavioral/developmental) and evolutionary supplies a focused paradigm in which proximate motivations are causal with evolutionary stable stratiges such that evolutionary theory can be constructed from the bottom (proximate mechancisms unconfounded with the paths of equilibrated games) up. The difference between the tournament and the display provides an outline in which the sensory biophysics to result so can be devolved into data processes in the process. Social selection does not reduce behavior to gene vehicles but supports a diverse set of concepts that can be empirically verified to see if true. Here I develop a particular social selection scheme for turtles via vestibular signaling by utilizing somatic programs that recapitulate prior unnecessary threat points and display a means that perfect teamwork telenomically in which coalitions of coincident self interest form providing a kind of cooperation in the social behavior of turles to rasie the most number of offspring in the next generation. As individually produced threats (to indivudal and competitive directions) create conditions for the discrimation of teams from the threat points per local interbreeding population somatic programs may constrain the mutations that can be beneficially incorporated. Decendent populations thus must keep fleshed in, older tissue supported threat directrums goaled towards cooperative functionality instead. Thus the somatic program houses computationally parallellized minimized payoffs that can not become independent of subsequent behavior no matter the negotiation or bargain.

One of the components of sexual selection is a consideration of secondary sexual characteristics. Here a particular physiological mechanism is proposed as a component of social selection that links selection of a sexually dimorphic trait exaptively to offspring emergent infrastructure through recapitulations of somatic programs developmentally. Behavioral strategies are reflected in the dimorphic phenotypes and differences in interpretion of the same traits based on sexual selection are contrasted. Mutual Sexual selection has been suggested for the head crest traits in non-avian Ornithodira (Hone et al 2012) The phenomenon of late appearance in ontogeny and profound ontogenetic change is taken as support for sexual selection of the trait but this will appear to be explained be social selection as well (formation of admission tickets AFTER coalitions form and need be costly enough to prevent non-earned entry). the idea that sexual display was their primary function has been fairly popular (e.g., Wellnofer 1991, Naish & Martill 2003, Unwin 2005, Bennett 2006). The fruitfulness of social selection is found in its providing a plurality of game theory phenotypes involved in optimizing life history strategies that underlies social infrastructure selection. Confusions about the theoretical and empirical boundaries of the process of social evolution are cleared up to some extent through the use of somatic program teleonomy to bridge between the two tiers in some cases Ernst Mayr developed the difference between proximate and ultimate causation in biology (from 1961) but some have challenged this distinction which relies for its truth on Pittendridgehs need to unconfound the immediate physiology from the evolutionary history. Social selection sensu Roughgarden with its implicit construction via two tiers (behavioral/developmental) and evolutionary supplies a focused paradigm in which proximate motivations are causal with evolutionary stable stratiges such that evolutionary theory can be constructed from the bottom (proximate mechancisms unconfounded with the paths of equilibrated games) up. Social selection does not reduce behavior to gene vehicles but supports a diverse set of concepts that can be empirically verified to see if true. Here I develop a particular social selection scheme for turtles via vestibular signaling by utilizing somatic programs that recapitulate prior unnecessary threat points and display a means that perfect teamwork telenomically in which coalitions of coincident self interest form providing a kind of cooperation in the social behavior of turles to rasie the most number of offspring in the next generation.

Potochink challenged Roughgarden to show that behavioral prefernences are strategic phenotypes within game theory kinematics. Here I develop a particular social selection scheme for turtles via vestibular signaling by utilizing somatic programs that recapitulate prior unnecessary threat points and display a means that perfect teamwork telenomically in which coalitions of coincident self interest form providing a kind of cooperation in the social behavior of turles to rasie the most number of offspring in the next generation. The quaternionic processing of the roughgardian reward sensation via the system of vestibular signallying sets up a signal receiver modality in wich acioms 1,3, and 4 apply in the strategic phenotypes trait wise available but variable through pareto optimality.

This is particular exemplified in the mormyrid fish adaptation of an typanmic air bladder and the modeling of the sound clicks as reflections and the moans as translations.

(Salamanders may have an internal market mechanicsm in pheromone production). The foreclaw display that uses the quternion belt-trick to divide out rotations performs behaviorally pleasure based locking by eliminating false torsion of different gibal systems associated with head-neck coordinate systems. This display is not found in side neck turtles because the needed motor patterns are utilized behaviorally differently. Head bobing in terrestrial turtles is also a part of behavioral interactions with the need to eliminate false gibal torsions during communication.

Suggestions for turtle sound evolution in general are generated from implications of C ( a smooth closed 3-manifold in H) and predictions different than sexual selection are provided by supposing social vestibular signal exchange of a certain natural quaternion-valued 3-forms relation to gravity via what was formerly called titillation.

There are two kinds of turtle foreclaw display behaviors: top-bottom and face to face (Krammer and Burghardt, 1998). Though these kinds of displays are generally divided between aquatic Emydid genera, juveniles have been observed to display both ways (Pseudymes (1998), Trachymes (personal observation)) . Juveniles have also been observed to display more often to objects than conspecifics which is the norm for adults. This behavior has been interpreted in terms of sexual selection as a means to facilitate forced intromission (Jackson 1972) but data to support this view have been lacking (Gibbons, SSD paper 2005?). It has been reinterpreted and renamed foreclaw display with an unknown communicative value (Thomas and Altig 2006)

It is suggested here that the goal of these different behaviors is manipulation of different neural processes from different parts of the 3 semi-circular canals and that this output is compared to what is heard through the inner ear through which programming by the belttrick across all the canals and macula enable turtles to play different individual or team firm incentives and thus the foreclaw display is divided into two different behavior computers with different affects on survival environments they co-exist in, by creating coalitions either at rest or during motion. These represent two different parts of optimal life history trajectories for aquatic emydids one of which returns in on itself and another in rectilinear motion. There is a uniform delay and acceleration between these two computers represented in the 1-D freize patterns sound environment that is permanent neubiologically distributed between pleasure and pain and thus persisting therein at the junction of the threat points and the NBS per NCE(some to the space and some to the velocity amongst translations and reflexions per rotations). Thus the social evolution of turtles depends on the difference of the composition of the velocities of cooperation and the spaces of competition roughly speaking. And so the proximate reward sensation enables a prevention of reduction to a narrower space in the NCE through cooperative NBS orbits (Closed) or trajectories (open) in the open somatic program.

The computations/algorithms from the belt-trick to untwist the rotations between translations and reflections outlines the paths game-theorized games equilibrate to, in the social selection of the infrastructure. Sound can be synthesized from actually infinite magnitudes of various frieze patterns distributed amongst the 3 canals as parts of the imaginary components of quaternions. Fan Scudder and Arial have shown that there is specific neuronal processing of just the lateral canals. The reflex of this action provides the reward sensation basis and locks the vestibular origin quaternion between two participants, providing a means to achieve teamwork as the participants climb their mutual but individually held pleasure gradients. Comparisons to sexual selection narratives are provided. The vestibular locking shows the falsity of trying a theoretical distinction between behavioral strategies and preferences (Alger). Locking sends the same information the brain as a strategy and a preference.
Parallelization in the Vestibular System: a differrent kind of parallel The biomechanics of the vestibular system is cognized relatively straight-forwardly. The doubled three roughly orthogonally oriented semi-circular canals provide information on head roll, pitch and yaw and the utricle provides help in determining linear acceleration and head tilt while the saccula (helps to determine relationships ) to gravity. Peterson, E.H. `1998 News Physiol. Sci. Volume 13 August 1998 Are There Parallel Channels in the Vestibular Nerve? Peterson (1998) attempted (to determine )if neuronal processing of vestibular system output follows that of other neurobiological structures wherein peripherally different parallel processes are delivered to the CNS through functionally organized groupings of inputs along the parallels. His analysis based on a review of four different shared characterizations in other identified neurobiologically parallel functionalities is generally negative for the vestibular system. The four properties found in other (systems) are: neuronal parameter heterogeneity, orderly parameter co-variation, grouping of neurons into statistically determined clusters even if parameter values may be bimodal or multimodal, and the variable cluster groups demonstrate functionally different sensory roles in the system of attributes (where different submodalities or subcategories of a stimulus possess qualitatively different information contents). Here a completely new vestibular biophysical functionality based on quaternions rather than euler angles is proposed and the possibility of a different kind of parallelization is suggested in the neuronal processing of vestibular system external input afferently sent through the vestibular system to the central nervous system.

Pitch, yaw and roll are the descriptors of Euler angles, a three angled system that can be used to describe generalized rotations in space. The computation of rotations using these angles however is not perfect and a phenomenon known as gimbal lock makes pefect computation of any rotation impossible and work-arounds are needed when applied to particular physical structures. Quaternions are an alternative mathematical representation for generalized rotations and physical instantiations of them do not suffer from gimbal lock difficulties. Computations with quaternions are however frustrated by the non-commutative nature of its multiplicative properties, which however is helpful in defining the intricate nature of somatic programs that cannot often be simply eliminated beyond vestigiality.

Euler angles The xyz (fixed) system is shown in blue, the XYZ (rotated) system is shown in red. The line of nodes, labeled N, is shown in green. So while the morphology of the semi-circular canals themselves will never experience gimbal lock (the xy and XY planes (fixed vs rotated) will never coincide morphologically) depending on how the processing of the angles is done in the brain (central nervous system) neural processes, may be put in a/the position to do a work around, as fluid moves through the semi-circular canals.

(or ) is the angle between the x-axis and the line of nodes. (or ) is the angle between the z-axis and the Z-axis. (or ) is the angle between the line of nodes and the X-axis.

Does the roughly orthogonal canal morphology express a fixed vs a rotatable state (wherein the macula output circumscribes the line of nodes or does each canal differently encode different euler angles at different times? (are there different parent hierarchies of the horizontal , posterior and anterior canals? (Does the common crus indicate that that the horizontal is hierarchically (because of gravity?) different than the anterior posterior??)) or could it present the 3 imaginary components of quaternions (with the macula indicating how unit quaternions are compounded)? Can one use a rational curve in a projective space or a rotation on unit quaterion??

From a practical point of view, a rotation of an angle vector

around an axis directed by a normalized

is represented by the quaternion

There is no problem similar to the gimbal lock with quaternions. This can be explained intuitively by the fact that a quaternion describes a rotation in one single move ("please turn radians around the axis driven by vector "), while the Euler angles are made of three successive rotations.
Does the macula normalize the parallel quaternion input from the 3 canals determined by difference is speed of pushpull pair deflections (slow fast) with neurons that fire regularly and irregularly? If hair cells can determine push-pull deflections (anterior posterior (two sides), left and right horizontal canal) with two different speeds, then the perpendicularity of the saccule utricle together can provide information to determine the operative unit quaternion and vestibular processing would occur as parallel data from the three canals is synthesized into a single quaternion.

Ernst Mayr has made what he considers a factually based case that Goulds attempt to explain the mechanistic relationship between ontogeny and phylogeny via heterchorony is incomplete. (1994) He describes somatic programs as recapitulations and challenges molecular biologists to find the ultimate why of his reasoning and explain how it is that some ontogenetic developments require somatic programs and others do not. These somatic programs were described as a behavior computer in 1961. Though some attempts to bring this idea forward have appeared (Digitial ontogeny/phylogeny; Measure of Teleonomic entropy) many comments have be negative (Hulswit, Ariew,Francis,Thompson).

I will show how social selection sensu Roughgarden is (dynamically)kinematically poised to differentiate the goal-directed processes implicated by the somatic program as behavior computers in the evolutionary tier by unconfounding the proximate reward sensation molecular biology from the behavioral equilibrium determining specific social infrastructures selected as adapted features by a direct application/instatiation/manifestation of the information metaphor Mayr introduced regardless of the teleomatic forces that the attraction and repulsion of the chemical bonds used/involved in necessitate. Thompson thinks that Pittenridghs idea that biologists who said, The turtle came ashore and laid its eggs rather than the turtle came ashore to lay its eggs meant to differentiate a description of an organization from an explanation of an organization. And that Mayr fails to do this. Mayr does say that proximate causes are purposive to the extent that computer programs are but that natural selection never is. Thus it will be possible to say that proximate reward sensation of turtles is TO have the most number of offsring and is done by the goal of either dipsplaying to the horizontal or the anterior posterior canals but because the molecular chemical bonds that that operate the program that does the goal direction may be many different kinds (4pie rotations in the dual electrical mechanical continuum)Plurals, the ultimate somatic program retained reacapitualtionally is not to be a particular adaptive feature even though particular adapted features do arise as particular chemical attractions and repulsions are historically accumulated (and expensive (effort needed to process increases in sound and vestibular amplitudes per frequencies) admission tickets in male claw length either do or do not arise). Thus Huxleys random small order variation that is converted by the operation of selection into directional large-order variation evolutionary change along lines adapted to the improved performance of biological functions in a given environment (1961) does exist in turtle organization. Because there can be different molecular materials that control the quaternionic monomial constants during the attainment of specifc goal directed ends, these ends nevertheless never are causally responsible for the current operation which is socially selected nevertheless. Social selection through specific somatic programs can be understood to fully unconfound the physiological mechanism from the evolutionary change.

Hulswit wrote, Mayr fails to explain what he means by causal. I can only interpret him as follows: a program is an efficient cause, which in combination with other efficient causes (the internal and external disturbances), completely determines the end state of the process or behavior. If so, then Mayr owes us an explanation of how under different circumstances, and thus, given different sets of efficient causes, the same program may lead processes to the same general end state. The behavioral tier equilibriums NBS under pefect teamwork generally do this.

More importantly: how can programs be considered as efficient causes at all? Or put more concisely: is there a theory of efficient causation that meets with the idea of a program? That seems unlikely, for efficient causes are singular events or facts, while programs are not. The different ways that the belt-trick can be employed topologically/algebraically and with what particular quantum mechanical rotations provide is a rich field for meeting this requirementans is extant. The singular quantum mechanical rotations may be in different chemicals but the 1-D freize pattern rotations are general for the particular interpretation of sound-vestibular functionality suggested here. There is also a possibility of using transfinite numbers of the patterns in the continuum (electrical-mechanical) should a better understanding of the relation of the telenomy and teleomatics was/be known.

Contrary to philosophical objections that Mayrs informational metaphor is outdated (Areiwe 2003) a particular somatic program of turtle vestibular signaling is proposed to underlie the proximate reward sensations that supply a pleasure gradient in which an individuals behavior computer exists somatically. The ultimate outcome of the proximate causation can be divided into teamworks, threats to revert to individualism, and back and forth negotiating, depending on the details of the proximate somatic program reality regardless of the logic. Sound discrimination (in contradistinction to the effect of the same sound on the vestibular system) provides the minimum intervals possible for back and forth bargaining. Mayr (1961) decribes a behavior computer of an individual as that that controls the development of the nervous system, sense organ, physiology and morphology as a whole. Roughgarden has shown how individualism and team work may have different trajectories in variable games theorectically. Mayr says, a behavior program that allows for appropriate learning and the improvement of behavior reactions by various types of feedbacks gives greater likely hood of survival than a program that lacks these properties. Which empirical programs are utilized can be determined experimentally and if supported falsify sexual selection. Application of Mayrs behavior program to social selection enables one to understand how he considered Ayalas use of teleology relative to the substantiated genetic code/genetic information as misinterpreted. Macneils dismissive of Mayrs innovation is likewise unfounded. http://evolutionlist.blogspot.com/2006/04/teleological-and-teleonomic-newer.html

The goals or programmatic directions are in the form of the equilibrium in the behavioral game and thus it is never the case that there is in nature a teleology to improve the reproductive fitness of a population through production of the DNA codes on the evolutionary tier since at

best there can be frequency variation/dependence/polymorphism (Roughgarden 2011) but not huge differences in the codes/genes themselves. Instead the behavior computer in the behavioral tier expresses only how they (individualism vs teamwork genes) are programmed behaviorally. The generality is that a population size can remain constant even if the behavior changes and the goal is different behaviors not different population sizes even though that could be possible. Different population size effects in this particular case would then be for differences between aquatic and terrestrial or semi-terrestrial emydids social evolution under probably a different social selection scheme. Later Mayr distinguishes this property of biological entities that appears not less purposive than the actions of a computer that has been programmed to respond appropriately to various inputs as teleonomy to distinguish it from teleology. This does not mean that teleonomic behavior computers that exhibit somatic programming are themselves purposive to the extent that human created computer programs are. There is a large difference between Aristotles use of the countably infinite and infinite divisibility. The specific predicative use of actual infinites in the synthesis of sound in the brain however can count divisions but only in a limited way (via the belt trick). The purpose man can put somatic programmed behavior computers to is severly overdetermined by the relations of recapitulation possible. Computer programs and cellular automata used to investigate these are not. Mayr introduced the notion of teleonomy and distinguished teleomatics from teleonomy sensu stricto. He even suggested that this philosophical dissection of teleology may function specifically in the call characteristics of birds. Here interlocking foreclaw pulsations vestibularly function instead. Mayrs use and reference to recapitulation have not been overwhelming supported but the introduction of social selection sensu Roughgarden may however find a subject to which this object may be causally connected as this view of the ritualized plesiomorphic foreclaw display is better understood as a pulsation of sound energy. Social selection with behavior programs enables the notion of prearranged information to be understood. Below is(largest amplitude dectetable smallest amplitude recoverable )

The combination of both may possibly be perceived by turtles by providing both right and left derivatives of quaternions and work by the Cauchy theorm from this boundary to an internal decision as to fire signaling receipt of the signal. The instructions that juveniles play out are different teleonomic entropy accumulations leading to different quaternionic monomial constants recoverable by transcendental identities within the same homogenous polynomial. This takes place neurologically beyond the vestibular nucleus which is still differentiating left and right canal outputs.

Here we say directly what it means to say that Mayrs program is material. We discuss how pleasure gradients are materially influenced differently by individual sounds and by the vestibular system as a reflex in teamwork through differences in type1 and type2 hair cells afferent outputs. Thus I determine the spatio-temporality of the Mayrian program in the quaterionic nervous processing of the Roughgardian reward sensation. It is possible to discuss explanatory and descriptive teleonomy within social selection. Teleonomic description depends

on just what 1-D symmetries are in the environments past and present of the teleonomy so being explained. The evolutionary tier is not preprogrammed but historical and depends on many things other than just the behavior computers churning while changing.
Goldberg and Harris suggest that the primary evolutionary transition in vertebrate crista is from longitudinally type2 only cells to centralization with both Type1 and Type 2 cells. If type 1 record translations and type 2 record reflections, the evolutionary transition would show a gain in ability to move the reflection origins. How can a reflection lead to a translation? Considering that the shape of the crista goes from latitudinal (fish-frog) to a torus split with a two central areas(turtle) to ovate longitudinal with a large central region and no torus, it may be that there was an interchange of vertical and horizontal reflections. A rotation caused by different neuronal connections (behaviorally mediated) to all three crista through a mutation may set up the possibility of gene fixation into a vertical reflection or simple translation or into both states (with a double faster vertical reflexion than the translation). This may have been what happened between anaminotes and turtles. A quaterinonic rotation may have been converted into a scalar fractal self similiarty. The evolution appears to present different spirals of quaternions.

If the canals each present a quaternionic axis then reversing the directions of these spirals is accomplishable by negating the axis. Is it possible that the push-pull (inhibition /excitation) in opposite side canals (left right/ anterior posterior)is a true negation of a quaternionic axis neurologically substantiated? As far as this goes, I don't think there's any one correct way to change the 'handedness' of a rotation in quaternion form. Negating the axis might work in some contexts, but in other contexts you may have to negate the angle (the w component), and also reflect the axis across a central plane. It just depends. Oh, and there's also the issue of what axis is considered to be 'up', which has nothing to do with handedness per se, but can also cause no end of confusion when importing models or animations. http://www.gamedev.net/topic/459925-quaternions-left-handed--right-handed/ The infinite frieze patterns may be different for the quaternionic imaginary parts and thus different for the canals as a whole thus permitting a rotation mutation to move into different homogneous quaternion set trajectory. This may occur by simple gene fixation behaviorarlly mediated. Combinations of different ordertypes of freize patterns per quaternionic system provides a construction from which sound itself can be decomposed into classes of 1-D symmetries (reflection/translations) rather than into fourier series. Consider each vestibular system to express a quaternion function (where a quaternion function is the same kind of thing as a complex function except that each canal gives the three imaginary parts and the macula preserves the scalar component). The left-right multiplicative dichotomy of sets of quaternions functioned thus is approached differently by anaminotes than reptiles and birds than mammals. This dichotomy is separated in time by fish and amphibians, by space amongst reptiles and birds and is involuted in mammals.

Each adapted system can be analyzed into the following concepts

holomorphy: existence of a difference quotient, harmonicity: satisfaction of Laplaces equation, analyticity: expressibilty in terms of power series, and conformality: local preservation of geometry

How each of these mathematical differences are divided in time, space and form provide a new basis to understand mutations of the vestibular system that may associate a simple rotation

translations

into a reflection that are conformal for instance.

that gives rise to

The difference between irregularly firing and regularly firing hair cells may express differences in analyticity. The difference quotient may be represented by the relation of the kinocilium to the coordinate space the quaternion is visualized (like in animations) in and the the urtricle, saccule perpendicularity may present in the striola a direct morphological element of the quotient. Morphological striola shape differences thus represent differences in how derivatives of quaternionic sets can be obtained in different species. Reflections and translations occur within these functions but the point difference may not infinitesimal in any case. The difference quotient ( f( x + h ) f( x ) ) / h could be replaced by a multiplication on the left and right by h http://www.zipcon.net/~swhite/docs/math/quaternions/analysis.html And though in pure math this buys us nothing in pure mathesis stoichiemetrically it already did. This occurs when a left and right behavioral impact (like left and right claws of turtles) effect a multiplication both on the left and right of the striola scalar effected left and right difference in all orientation afferent signals of the hair cells. Thus the nervous system can do better than the mathematician in being able to use this means as well decomposing the sets from heterogeneity to homogeneity within the form only of functions of x for which the limit of a right difference quotient exist are of the form p + qx, and those for which the limit of a left difference quotient exist are of the form p + xq because the vestibular system (WITH SOUND) can find differences in 1-D symmetry (translations or reflections) WITHIN the difference of these two mathematical forms limited (approaching from the right or approaching from the left). P and X have different frieze patterns with respect to q. These differences can be mathematically described by using different actually infinite numbers of the freize patterns in each imaginary part of Q while the translation difference is variable in the scalar.

An infinite strip with a symmetric pattern is called a frieze pattern. There are only seven possible frieze patterns if we are using only one color. Note 3 1. Translation symmetry only:

2. Glide reflection plus translation symmetry:

3. Reflection over a horizontal line plus translation:

4. Reflection over a vertical line plus translation:

5. Rotation (a half-turn about a point on the midline of the strip) plus translation:

6. Reflection over a vertical line plus a reflection over a horizontal line plus translation:

7. Reflection over a vertical line plus glide reflection plus translation:

The infinite strips may be of different cardinality and ordinality per ordertypes. And thus biology provides math with a practical system to investigate the actually infinite. The physics of force application (Pseudymes from top to anterior posterior Q axes with left and right hands (two axes), and other Emydids to left and right by left and right hands to horizontal imaginary Q axis (one axis)within this structure is the first instance of application of transfinite numbers to physics.

Turtle Titillation: Can turtles hear the foreclaw display?

Many Emydid turtles perform a seemingly rather unusual behavior, originally termed titillation. One turtle approaches the other, either from the top or in front, stretches its forelegs towards the other and vibrates both front feet, often contacting the ocular region or the carapace of the conspecific.

Numerous videos on YouTube demonstrate some of the variability in this behavior, as well as, the nonspecific teleonomic nature of its object. Turtles can be seen to vibrate or titillate at fingers outside the tank as well as towards, nonmoving objects within.

http://www.youtube.com/watch?v=AmY0cFv3was&feature=related(1 picture) nd http://www.youtube.com/watch?v=k-Jay_tZLCo (2 picture) (another video directed at an animated object below) http://www.youtube.com/watch?feature=endscreen&NR=1&v=XkiOO114LqU http://www.youtube.com/watch?v=_KUatkmXp3o Recent studies (listed in Thomas and Altig 2006, pictured in Karen Davis 2009 below) have shown that the activity is not restricted to males targeting females as was traditionally described but is found amongst juveniles, across species, within the same sex, by females onto males, and by putative dominants to ostensive subordinates. Due to this diversity of aspect in the context of the unknown nature of the communicative value of this interactivity, Thomas and Altig 2006 have suggested renaming it, foreclaw display.

st

But is it a display? Is it all about the eyes or is it a sound-energy-transfer-pulsation directed at the ears that may be pleasurable as well providing the means towards mutual direct benefit ends?

I have kept a small male Trachemys scripta with an adult female Chrysemys picta for the past 5 years.

When the male was a little larger than the size in the photo above (here~60 mm), he began to vibrate his foreclaws at the female by swimming on the top side of her (with shorter finger nails this may be an easier way to reach the ear), as occurs normally in Pseudemys (Citation ?). Now he is much larger and only displays from the front, as is common for both genera (notice the female declines her head by propping up her hind legs(below)). The male has been observed to produce up to 52 consecutive drumming bouts in one series. Although most of the sequences are by the male scripta onto the female picta, the female has been observed to vibrate her forelimbs onto the male as well. Female Chrysemys picta titillation has been observed (Ernst 1971) but not female Chrysemys titillation onto Trachemys? Rives 1978 (I have not read this yet) has investigated titillation between these two genera.

Last year, the pair mated (a new observation for these two genera) but did not produce viable eggs (no embryo spot could be identified in any of the eggs). Some of the eggs had extra calcium bulbs at their ends. Examples of cross genera foreclaw pulsation interactions:

Davis2009 The sound of the foreclaw drumming is so loud, usually impinging on the carapace, that it can easily be recorded outside the tank, with a simple entertainment microphone (that came packaged with a Dell Inspirion mini-10). Sonic Visualiser software (http://www.sonicvisualiser.org/) indicates that the frequency range of the sound is between 1,000 and 6,000 HZ and the average frequency of the claw vibration at about 20 HZ per bout.

(I probably have about 150 of these traces to quantify(intra and inter claw drumming temporalities) Can this sound be heard by the turtle. The Eqyptian tortoise has a very similar sound and it seems obvious to assume that these turtles do hear them. The head moves very easily in water and I propose that the foreclaw pulses energy into the heads that can be detected by the vestibular system and heard by the inner ear

Thus the display which begins with a visual posturing of the forelimbs may not simply be an ornamental sexually dimorphic trait subject to runaway sexual selection rather it could be a proximate behavioral reward sensation, causal to social infrastructure selected? (Roughgarden et al 2006 2010? 2011 ? 2012). A selective pressure which may serve in the evolutionary tier as individual fitness is behaviorally accumulated may be derived from the following figure:

What is the function of the behavioral communication and wherein does its social context remain. What is the meaning of the potentially plesiomorphic ritualized juvenile behavior and what is the behavioral evolution of this trait. Must it being generally sexually dimorphic necessarily be cast in terms of intra vs inter sexual selection? What are the senses involved and how often and under what contexts does it appear in nature?

Manley reports that Preferred intervals in the spontaneous activity of primary auditory neurons, a probable correlate of electrical tuning, have been reported from turtles, lizards, and birds (25). (2000)

in Comparative Hearing: Birds and Reptiles, Springer Handbook of Auditory Research, eds Dooling R, Popper A N, Fay R R (Springer, New York) , in press. Can the frequency of the foreclaw pulsation be detected by auditory neurons? And could these neurons signal pleasure?
The intensity of the foreclaw display (as shown above) drops off to the background level below 1kHZ so the turtles may be deaf to the frequencies of the display itself but perhaps able to sense the 20 HZ bout as a whole, either through a tuned cell to the foreclaw display biomechanical frequency (literature indicates tuned cells either down to 10, 20 or 30 HZ) directly or perhaps through some kind of mechanical-electrical resonance and or via sensitization from afferent vestibular signalling. (This difference would affect the physiology of the reward sensation motivation and the empirical nature of the equilbria/games structurable. ) Single pulse impacts of sound or voltage have oscillating decreases through 3 cycles at the tuned frequency (Crawford and Fettiplace)(illustration below). Interestingly hair cells divide into two types (Art and Goodman) below and above 60HZ (2 * 20) thus a A 20 HZ foreclaw display may thus possibly resonant with a proximate biophysical situation where changes in channel types occurs. This provides a rich field for the structure of the games that can formatted under theory. Could it be that the 20Hz sensitivity provided the structures later evolved into sound production by Testudines? Inner ear hair cells are not passive reporters of mechanical input (Art and Goodman 1996). Here I suggest that the input is mechanical sound energy and that turtle titillation is a communication medium of mechanical sound pulsations. Frequency tuning occurs after the transduction of mechanical

energy(Art and Goodman page 106) thus the biomechanics of behaviorally gamed forelimb muscle motion creates energy that can be tuned to purely electrical energy of the same waveform and forms the basis for learning and reciprocity over evolutionary time.

(This needs to be reillustrated to reflect 20 HZ 60 HZ resonance not 10-30HZ and done in a different manner) Closing of the eyes may put pressure on the inner ear itself, perhaps resetting the vestibular system.

The trait is probably much more common, variable and diverse than has been documented to date and due to the long periods of time captive specimens can be seen to devote to this behavior, surprisingly, the full functionality of its behavioral ecology remains largely unelaborated. The senses involved in the activity have not even been quantified. The sound frequency distribution of the foreclaw display is very similar in many respects to the sound made by the Epytpian tortoise in close physical intimacy. Can it be that The foreclaw display is heard rather than simply seen?

Can the 60Hz differentiation of cell types be associated with fitness accumulations on repeated behavioral bouts of foreclaw pulsations socially selected. Can NBS teamwork have caused the diversification of cell types above 60 HZ epigenetically? Can the foreclaw display energy impacts prevent zeroing the vestibular system causing teamwork instead and thus inhibiting asymmetrical reversion to individualistic threat points ? Do somatic programs ontogentically constrain the evolution of trait associations or is this system a runaway good genes structure in the population. Painted turtles posses a neural equivalent of eye-blink reflex in which electrical stimulation within the brain can accomplish the equivalent of muscular contraction of the eyes. It may be that the foreclaw display is alternatively directed at this physiology if the extraocular nerve firings only can effect electrical changes in other head nerves with sensory elements (thus permitting the foreclaw display to be part of the a general head sensor of others or if the direction of conduction can be reversed (from the eyes to the vestibular or ears to the brain) of the ear or works in conjunction with hearing.

With this preparation, a neural discharge can be recorded from the abducens nerve, which projects to muscles controlling the extraocular muscles, after a single shock electrical stimulus is applied to the ipsilateral trigeminal nerve [the unconditioned stimulus (US)] that contains sensory fibers from the

head. This discharge represents a neural correlate of the turtle eye blink reflex (Keifer, 1993a). When this stimulus is paired with a train of stimulus pulses applied to the posterior eighth, or auditory, nerve [the conditioned stimulus (CS)], after a period of time a positive acquisition slope of CRs can be measured in the abducens nerve (Keifer et al., 1995). After unpaired stimuli such as an alternate-pairing training protocol in which the CS precedes the US by 10 sec, the CRs extinguish and show reacquisition when pairing is resumed.

As this is it seems that the foreclaw display through sound combined with tactile sense may cause the eye to stay closed as a reflex.

Title Index

Book Series

Non-serial Titles

Book News

Brain, Behavior and Evolution 1998, Vol. 51, No. 4 Central Trigeminal and Posterior Eighth Nerve Projections in the Turtle Chrysemys picta Studied in vitro Herrick JL, Keifer J

It may be that the turtle only feels the contacts of the foreclaws on the skin or the eyes (and thus offer access to a commom pleasure gradient neurologically) , but given the intensity of the sound energy (some measure of this should be included) and the characteristics (and coincidence of properties ) of the inner ear cells to physical deformation, it seems likely that the cochlear or the vestibular cells or both, are the physiological concomitants of the sensation. The turtles can thus feel the pleasure of the other directly by the sensitivity of their hair cells to different incoming frequencies because the hair cell can both effect mechanical output and move with incoming input. Both turtles are on the same page as they play as juveniles and establish coalitions with similarly responding populations of hair cells to slightly different frequency profiles. Trachymes audio show differences comparable to those sound differences selectively chosen by female Testudines but are the sounds produced and the potential sound reception of the foreclaw pulsation causally related? How well can they be correlated?? The brains evoked potential though preferring 100-400 HZ can receive ossilations from 40 to 6,000HZ. Is it true that turtles just dont use sound or rather is it otherwise?

( The behavioral evolution of this trait remains to be determined. Aspects includea apparent phylogenetic variation with possible ritualized plesiomorphic ontogenetic expression(Kramer and Burghardt 1978, reviewed in Burghardt 2005)). If the ears are the target organ for the behavior then the selection pressure for longer nails may be indicated from the following: There are differences in the position of the turtle, the length of the nails, the time of the year, the association with copulation, relations to the posture, number of bouts during a series, sequences of other interactive behaviors. Does biting always indicate an agnostic action. Could biting effect the vestibular system by causing whole body shocks and thus be related to pleasure in some way?

This of course only takes into account the nail length and not the motor pattern itself nor the display posture that precedes the act.

I Cochlear mechanisms from a phylogenetic viewpoint 1. Geoffrey A. Manley*

PNAS October 24, 2000 vol. 97 no. 22 11736-11743

Preferred intervals in the spontaneous activity of primary auditory neurons, a probable correlate of electrical tuning, have been reported from turtles, lizards, and birds (25).

1. 25-Gleich O, 2. Manley G A (2000) in Comparative Hearing: Birds and Reptiles, Springer Handbook of Auditory Research, eds Dooling R, Popper A N, Fay R R (Springer, New York) , in press.
Are the auditory neurons preferring / able to prefer intervals originating from foreclaw pulsations?
The efferent connections of the nucleus accumbens in the lizard ... www.springerlink.com/index/t46837t52444225h.pdf by WJAJ Smeets - 1995 - Cited by 28 - Related articles nucleus accumbens and, caudally, to the peribrachial region and the superior ..... whereas in turtles a nucleus accumbens is hardly identi- fiable (Smeets 1988).

Roughgarden has written (2012)

It is hypothesized that natural selection would cause the evolution of teamwork instead of individualism because teamwork would be beneficial to the individual on average. Imagine a gene for individualism and another gene for teamwork. The first codes for pleasure sensory neurons that respond to an increase in one's individual fitness increment irrespective of the effect on others. The second codes for pleasure sensory neurons that respond to an increase in one's

own fitness increment as well as to an increase in the fitness increment of another that is communicated to it through pleasurable physical and/or vocal intimate interactions. The carrier of the gene for individualism might sometimes be better off because of no urge to compromise, but the individualism gene might also find itself in a role where it would be worse off than a bird who could compromise and work in teams. Given that the carriers of a gene will face an ensemble of different situations, it is hypothesized that the individual who works well in teams will, on the average, accrue a higher lifetime fitness than the individual who is individualistic (Roughgarden, 2009, Chapter 8).
Intracellular recordings were obtained from brain slice preparation in neurons of the striatum of the turtle Trachemys scripta elegans, analogous to the mammalian striatum in its topographic organization, synaptic connectivity, cytoarchitecture, and neurochemistry. Here we show that these similarities extend to the electrophysiological properties of its neurons. Biocytin staining revealed that 85% of the recorded neurons were medium spiny neurons while 15% were aspiny neurons. Spiny neurons of the turtle resembled those found in the mammalian and avian striatum and express dopaminergic D1 and D2 class receptors. Because the striatum of the turtle receives a dense dopaminergic innervation from tegmental dopaminergic neurons we investigated the postsynaptic actions of selective dopamine receptor agonists in the excitability of spiny neurons. As in mammals and birds, activation of D1receptors enhances, whereas activation of D2-receptors decreases the evoked discharge. Apparently, actions of dopamine agonists occur via the modulation of L-type (CaV1) Ca2+-conductances. Strong cellular evidence suggests that the role of dopamine in the modulation of motor networks is preserved along vertebrate evolution.
Cellular and Molecular Neurobiology Volume 30, Number 5 (2010), 743-750, DOI: 10.1007/s10571-010-9499-7 Original Research Dopaminergic Modulation of Spiny Neurons in the Turtle Striatum Jaime Barral, Elvira Galarraga, Dagoberto Tapia, Edn Flores-Barrera, Arturo Reyes and Jos Bargas

Dopamine and pleasure centers -

If the selection pressure on the foreclaw length is associated with increases in sound energy communication into ears then the functional physiology of individually derived fitness increases need to not develop into an anatomical system capable of combining the fitness increments of others. Since ear hair cells posses both efferent and afferent fibers it is possible that pleasure sensory neurons may efferently multiply teamwise what otherwise are individually based pleasures through the mechanical electrical amplifer. The pleasure which comes in from a mechanical (intimacy) creates pleasures which are electrically amplified either based on individual selfish outputs or via means of theteam. NE threat points may be realized within the nature of the hair cell amplifer itself.

Efferent synapses can alter cellularly fixed resonances and thus input from the eyes or skin in addition to afferent forcelaw produced electrical ossilastion nervously signaled may permit closer behavioral interactivity and changes in roles presented evolutionarily. The NBS NE could be encoded by the two way activity of the electrical tuning (afferent efferent) but only a one way effect of the mechanically forced tuning (throught the hair cell). There would be not fallacy between cooperation and competition as suggested by Ghiseln but simply different molecular means to accomplish signal amplifications. Crawford and Fettiplace (1980) have suggested that the auditory range reflected through the distribution of the characteristic frequencies of isolated cochlear hair cells of Trachemys scripta is 30 700HZ. Sea turtles may be sensitive to higher frequencies and the frequencies of Chelodina seem to suggest higher ones as well. (How to explain this?) Fettiplace and Fuchs suggest that tuning is no possible above 1 KHZ but if as is suggested below their may be a social infrastructure deme effect higher frequencies may have been tuned by the mechanical properties of lower order motions into sounds that overgenerations changes the molecular capabilities through the common mechanical electrical resonance where multiplicative tuning is done by team work accomplished mechanically but feedback electrically. Thus turtles unlike frogs or lizards are able to accomplish higher frequency (above 1 kHZ) selectivity by electrical mechanical cybnertics rather than through different kinds of pre-mechanical tuning.

The intensity of the foreclaw display (as shown above) drops off to the background level below 1kHZ so the turtles may be deaf to the frequencies of the display itself but perhaps able to sense the 20 HZ bout as a whole, either through a tuned cell to the foreclaw display biomechanical frequency (literature indicates tuned cells either down to 10, 20 or 30 HZ) directly or perhaps through some kind of mechanical-electrical resonance. This difference would affect the physiology of the reward sensation motivation and the empirical nature of the equilbria/games structurable. Single pulse impacts of sound or voltage have oscillating decreases through 3 cycles at the tuned frequency (Crawford and Fettiplace)(illustration below). Interestingly hair cells divide into two types (Art and Goodman) below and above 60HZ (2 * 20) thus a A 20 HZ foreclaw display may thus possibly resonant with a proximate biophysical situation where changes in channel types occurs. This provides a rich field for the structure of the games that can formatted under theory. Could it be that the 20Hz sensitivity provided the structures later evolved into sound production by Testudines? Inner ear hair cells are not passive reporters of mechanical input (Art and Goodman 1996). Here I suggest that the input is mechanical sound energy and that turtle titillation is a communication medium of mechanical sound pulsations. Frequency tuning occurs after the transduction of mechanical energy(Art and Goodman page 106) thus the biomechanics of behaviorally gamed forelimb muscle motion creates energy that can be tuned to purely electrical energy of the same waveform and forms the basis for learning and reciprocity over evolutionary time.

(This needs to be reillustrated to reflect 20 HZ 60 HZ resonance not 10-30HZ and done in a different manner) Closing of the eyes may put pressure on the inner ear itself, perhaps resetting the vestibular system.

Julian Davis A computational study into the effect of structure and orientation of the red eared slider utricle on hair bundle stimuls Silber Analysis of vestibular hair cell bundle mechanics It may be that the turtle only feels the contacts of the foreclaws on the skin or the eyes (and thus offer access to a commom pleasure gradient neurologically) , but given the intensity of the sound energy (some measure of this should be included) and the characteristics (and coincidence of properties ) of the inner ear cells to physical deformation, it seems likely that the cochlear or the vestibular cells or both, are the physiological concomitants of the sensation. The turtles can thus feel the pleasure of the other directly by the sensitivity of their hair cells to different incoming frequencies because the hair

cell can both effect mechanical output and move with incoming input. Both turtles are on the same page as they play as juveniles and establish coalitions with similarly responding populations of hair cells to slightly different frequency profiles. Speculationally, it may be, this may happen, that the afferent foreclaw mechanical input, say to cochlear hair cells results in efferent electrical oscillation biases in other hair cell populations ( perhaps the vestibular) such that the behavior results in a truly reciprocal interactivity that has reflex like effect adjusting the recipients sensitivity to its ability to sense and negotiate 3-D space depending on the length and during of the bout and giving rise immediately to changes in relative physical spacing possibilities in the habitat conducted physiologically. The extra length of the foreclaw may be analogous to the snake quadrate and/or exist as a form of a motor training hypothesis (Burghgardt 1998) wherein motor patterns connect and reconnect synapses with a reward sensation component. Head bobbing in Testudinates may be head not leg vestibular to inertial responses instead. This interpretation supports the idea of how the interactivity effects a direct negotiation through bargaining without language in the 3-D water environment(suggesting implications for evolution of the turtle hearing as due to play that leads either to adult titillation differences and to sound communication modalities (without a strict difference of only agonistic and sexual ethological categorizations). In cases of picta scripta titillation Rives 1978 has suggested that the display is aversive to the recipient and while this may have appeared the case to me early on when the pair first started the behavior, as the male grew in size over time it appears that the female stopped trying to avoid the male (by putting her head in a corner and thus being unavailable) but began to seemingly willingly drop her head and allow the titillation to continue for long periods, ending sometimes only if/when the claws projected inside the carapace. Whether it is social dominance or simply coalitional formation is difficult to assess. The divergent categorization of agonistic vs sexual may not be all of the ethologies involved here. Regardless the possibility of perfect teamwork is embodied in the biophysics of the electricalmechanical double continuum of the hair cell biochemistry and anatomy. The cochlear cells have been divided into two kinetic regimes that differ for low and high frequency. Low frequencies up to 60HZ are found in KV cell types and in over 60HZ BK cell types are extant. Propositionally, within a group of cells it may be a case that turtles have used the behavioral forelimb motor patterns to effect a somatic environment subject to somatic programming (Mayr) that trades-in, the temporal regularity of the motor pattern at low frequency KV Potassium channels for changes in sensitivity and frequency discrimination in K+ BK channels epigentically. In other words fast behavioral NBS-NE (Roughgarden et al) could result in different RNA splicing situations passed onto other hair cells after attainment/entrainment of a behavioral game and this would be subject to genetic polymorphisms over evolutionary time. This is highly speculative but the possibility is open for aquatic turtle ancestors to have created aptive somatic conditions later evolved in other populations for sound communication both on land (Testudinidae) and under water Chelodina oblonga as slow rectifier K+ currents are used to stabilize higher frequency populations (upto 200HZ)??? Turtle hair cells adapt to repeated deflections of the hair bundle (which if the foreclaw display is a sound pulsation would be doing so) by Ca+ dependent negative feedback on the opening probability of

the transduction channels. Can the behavioral trade-in of motor synchrony proximately effect general effects through hair cell adaptation? Can Ca+ binding affect other general tuning actions in concert (accessory structures, voltage-dependent dependent properties, and afferent transmitter release) If one turtle gets pleasure from the pulsation can may remain being pulsed long enough for afferent transmitter release to be effected through adaptation itself rather than through tuned resonance? Since efferent shocks to hair cells decrease sensitivity especially in low frequencies to sound input,
November 1, 1984 The Journal of Physiology, 356, 525-550. Synaptic hyperpolarization and inhibition of turtle cochlear hair cells. 1. 2. 3. J J Art, R Fettiplace and P A Fuchs

pleasure may provide a reward sensation directly this way requiring more pulsations for continuing the same amount of climbing the pleasure gradient. Thesis -Side-payments and teamwork may be differentiated by the relative amounts of electrical and mechanical contributions to the tuning of cells (teamwork electrical , side-payments mechanical) Pleasure based teamwork may function through alternative slo splicing variant expression whereas sidepayments may arise by changes in the mechanical movement of the hair cell. recent provocative studies point to a key function of chromatin structure and histone modifications in alternative splicing regulation. These insights suggest that epigenetic regulation not only determines what parts of the genome are expressed, but also how they are spliced Can turtle foreclaw pulsation coalitions create different epigentically controlled splice categories? Alternative splicing may be a basis for somatic programming telemactivally causative in the NBS NE social dynamics. The threat points (reversion to individual (beyond even side-payments) may be directed by the ability to cause histone methlyation on behavorially equilibrated foreclaw pulsations. The reversion to individuistically accomplished threat may be effected by reliance on the mechanical rather than the electrical contributions to the continuum access to pleasure based neurology. Thus a difference between perfect teamwork and various forms of partial cooperation would depend on how the two directions of the electrical continuum related to methlyation exposed histone splice variants during juvenile to mature growth/ development. It has been previously assumed that sexual selection drives the evolution of all the kinds of turtle titillation (Berry and Shine 1980 ) properties but it may also be possible that if the ear is the object of turtle titillation, that turtle play (found for instance in softshell turtles ,Burghardt) resulted in behaviorally capitulated evolution of hair cell tuning, through foreclaw pulsations programmed somatically and selected socially. It is possible that behavioral dynamics equilibrate epigenetically determined heritabilities of K+ channel differences in different social infrastructures. Different social group structures may result in different epigenetic variations that are ESS where no mutant can invade (without destroying the trait itself). Can it be that as new hair cells are histologically differentiated that

use and disuse of the foreclaw display effects retained RNA splice variants that simply accumulate as the turtle ages and that this aggregation of effects combined with group coalitional formation results in more offspring produced without any competition per say but by cooperation behaviorally to demonstrate the foreclaw pulsation. Fettiplace and Fuchs(1999 Ann Rev of Physiology) reviewed that , the BK channel which is thought to found the electrical tunning in the higher frequencies, is most likely the product of a single slo gene originally described in Drosophila and later found in the mammalian brain. They suggest that since multiple alternative splice sites exist and that some of these splice variants posses different channel kinetics that variation in tuning frequency can be correlated to differential expression of the splice variants. Only two of the potential 8 splice sites are used by the turtle. Can social selection through epigenetic cell memory by changes in game matrix payoffs in evolutionary time be related to the (existence and) selection of the splice sites themselves? How is this biophysically possible? Fettiplace and Fuchs note ion channel expression may be related to the synaptic organization of the hair cells. It may be that the mathematical topology of the synaptic connections which is determined by the actually developing (from juvenile to adult) nervous system (electrical-mechanical double continuum) constrains the possible genetic quantities available to selection. Can the synaptic organization and the reward sensations telenomically be in cause and effect with multiple slo gene accetylations over generations? Can multiple accetylations end up telematically (Mayr difference of teleonomc and teleomatic) through Nash barganining equilibria (over individual NE) to telenomically target specific histone areas of the slo gene resulting in specific slice cite variants being systematically exposed?

http://w3.biosci.utexas.edu/atkinson/atkinson/Projects_Epigenetic_Reg_slo.html

Interestingly in Trachemys scripta there is a transition towards whom the foreclaw vibration is directed at as the turtle ages (Thomas 2002). Young males often pulsate towards females but older melanistic males do not but tend to orient their activity on average towards (younger) males instead.

Does this reflect a difference in sexual selection or social selection? In the case of social selection it could simply be that turtles are able to effect more offspring in the next generation by associating in coalitions of foreclaw pulsing individuals since older melanistic males may be indicators or badges of more years reproducing and that it is female site specific growth that is correlated with age of male melanism (discussed in Thomas 2002). So female choice may not simply be selecting for males with good genes as symbolized in foreclaw length but instead ranked coalitions of foreclaw pulsing individuals behaviorally equilibrated may effect more offspring in the next generation simply ontogenetically and behaviorally as teams and this evolution, may have resulted in the diversity of auditory sense in chelonians through recapitulation (Mayrs new sense of) of developmental pathways. With male melanism being a resource holding potential observable badge to whom younger turtles not in coalitions would tend to attempt to associate with. The view proposed here, that turtles titillate/ demonstrate the foreclaw pulsation through the medium of the inner ear offers a potential means to identify the/those pathways opened up behaviorally and constrained physiologically. Gibbons and Lovhich suggest that with respect to sexually dimorphic traits that natural selection should operate equally on both sexes while they are juveniles with similar sizes and behaviors, i.e. prior to attainment of maturity. If turtle titillation is a complex auralelectrical feedthrough behavioral ecological structure forming trait,
Bidirectional transduction in vertebrate hair cells: A mechanism for coupling mechanical and electrical processes

T.F. Weiss

a, b

Hearing Research

Volume 7, Issue 3, August 1982, Pages 353360

It has been proposed that the sharp frequency selectivity exhibited by cochlear hair cells and neurons of alligator lizards results from a mechanical resonance of the stereociliary-tectorial structures of hair cells. In contrast, in the red-eared turtle this selectivity has been attributed to an electrical resonance mechanism located in the hair-cell membrane. In this report a new mechanism is proposed, one which is consistent with observations in the lizard and turtle preparations and in which hair-cell resonances result from coupling of mechanical properties of hair-cell stereociliary-tectorial structures with electrical properties of the hair-cell membrane through a receptor membrane process that has bidirectional, mechanoelectric and electromechanical, transduction properties. This same mechanism could also be the physical basis for diverse phenomena observed in mammals, including changes in mechanical properties

of the ear in response to electrical stimulation in the cochlea and central nervous system, and the presence of sustained, narrow-band, acoustic emissions in the ear canals of humans.

then this assumption which lies behind how to categorize the difference of sexual and natural selection in turtles may not be operative nor realistic. The view proposed here indicates that foreclaw pulsations can provide a bargaining leverage even after the older melanistic individuals die. Thus through the details of this synthesis one could empirically approach if Trachymes tend towards individualistic payoff plays vs joint working to attain the NBS (as considered in correcting Binmore (Roughgarden 2012) The question remains if the inner ear hair cells can differentiate with or without association with neurons.
Actin Filaments, Stereocilia, and Hair Cells: How Cells Count and Measure Annual Review of Cell Biology Vol. 8: 257-274 (Volume publication date November 1992) DOI: 10.1146/annurev.cb.08.110192.001353 L G Tilney, M S Tilney, and D J DeRosier

What is the hair cell transduction channel? 1. David P. Corey

1

October 1, 2006 The Journal of Physiology, 576, 23-28. At the present time we do not know the molecular identity of the transduction channel, but these

previous apparitions are bringing us closer to an answer. We know very well what the hair cell transduction channel should look like, in physiological terms. It is a non-selective cation channel that is permeable to all the alkali cations and to many divalent cations (Corey & Hudspeth, 1979; Ohmori, 1985). The channel has a particularly high calcium permeability, passing Ca2+ at least 10 times better than Na+ (Lumpkin et al. 1997; Ricci & Fettiplace, 1998); at the same time, Ca2+ in millimolar concentrations acts as a partial blocker, inhibiting current by monovalent ions. Remarkably, the channel is also permeable to small organic cations, even fluorescent styryl dyes like FM1-43 of molecular weight > 450 Da (Corey & Hudspeth, 1979; Gale et al. 2001; Meyers et al. 2003; Farris et al. 2004). There are no highly specific blockers: amiloride and its analogues, as well as aminoglycosides, block at low micromolar concentrations in a voltage-dependent manner. The block is relieved at more negative potentials (Jorgensen & Ohmori, 1988; Kroese et al. 1989; Rusch et al. 1994) suggesting that these charged compounds can be driven through the channel by voltage (Marcotti et al. 2005). A systematic study of blockers indicated that the pore is 1.3 nm in diameter at its narrowest, with a larger vestibule extending 1.5 nm in from the outside (Farris et al. 2004). Perhaps consistent with the lack of selectivity, the conductance of the channel is large, ranging from 100 to 300 pS or more

(Crawford et al. 1991; Geleoc et al. 1997; Ricci et al. 2003). That the conductance can vary by 3-fold or more in different hair cells from a single organ (Ricci et al. 2003) was quite unexpected, but it puts further constraints on prospective candidates.

If sexual selection is operative and males and females both benefit from maturity at an earlier age then on that view, turtles would be sexually selected to enter the breeding population earlier since this allows them to mate more often and thus have individually higher probabilites of successful breeding. On an alternative social selection view, turtles are selected to join social coalitions of older age founded titillating individuals, within which groups females have titillated within when younger, thus resulting in a larger probability of offspring surviving into the next generation than other titillating groups that may have founded around less melanistically badged individuals. One view leads to selection for particular genes and the other for social infrastructure. One says that maturation at small size is evolutionarily stable while the other that a myopic proximate mechanism finds a global objective correlated with larger size and is thusly stable. So here we can see that in two particularly specific example hypotheses of sexual and social selection in Trachemys, implicate opposite extremes of morphological extent (in growth). Nesse has asserted that social selection or social infrastructure selection (Rougharden, Oishi, Ackay) is confused and that it is a superordinate set over sexual selection as a subtype logically and that it is not an alternative . This claim is only a semblance of the truth as it confounds or confuses itself logic in theory and realizations in practice by not addressing the identities needed conceptually between theory and pratice and thus substitutes a probability In the application for a versimultude between any possible hypothetically developed alternative no matter the differences in theories themselves. The possibility that turtle sizes in populations may be explained by sexual trait selection or social infrastructure selection does not mean that the sexual selection supported traits need be within the social infrastructure set wise. If the social selection view is wrong in this hypothesis presented there may still be infrastrucutures being selected but it may be there is a good enough gene able to be runaway selected and a means to generate bad genes. Gibbons and Lovich suggest that natural selection seems to be operative for turtles smaller than 90-100 mm plastron length, since maturity is rarely attained by males below this range, regardless of age. Vestibular hair cells increase in number with size of the turtle. It may not be that mating is the clinamen between sexual and natural selection. The size associated threshold may be caused by learning and play in juveniles that appears not with age but with size as more tissue is added and more affected cells become part of an acting organization. The assumption that natural selection (social selection) operates in a egalitarian way amongst juveniles could be false. Thus it is not maturity and post mating action with active competition that is operative but ontogenetic irreversibilities (which includes maturation also) caused by juvenile activity prior to maturation and cooperative game theory equilibria projected beyond (responsible for changes to higher frequency sensibilities) that is. Below 90-100 mm, turtles simply do not have enough hair cells for ritualized titillation to have its full physiological effect. It is definitely possible for turtles to form the equivalent of binding contracts neurologically and thus bargain without the use of language if this suggestion is borne out and Gibbions and Lovich s dissection of natural and sexual selection is mistaken. Grand coalitions (all individual is a deme or population) would be

precluded from forming in turtles due to the directional information asymmetries carried mechanically and electrically. The non-adaptive nature of the sexually selective traits depends on what the sense mechanism is that controls the relationship between social infrastructure and the relevant behavior however expressed. Soc

If the changes in the number of hair cells in the Ultra macula is indicative of inner ear hair cells generally the change from hatchling to medium sized turtles (4,000-8,000) vs 12,000 in mature large sized turtles is crucial for the behavioral ecology. Newly generated hair cells have small surface areas and thin stereovilli. So it may be that the fast behavioral dynamic proximate reward sensation playe by juveniles architecturally format the thickening and increases in surface areas of the hair cells. Growth is not always related to age. Growth is highest just after turtles hatch (25-30% increase can occur in short time). Growth in the UM seems to decrease asymptotically as does growth in general at maturity. Turtle UM hair cell densities are constant over time which is different than found so far for amphibians and birds. The kinocilium of small cells are often short resembling the other ciliary rods. The constancy of the foreclaw display frequency may operate to adjust the height of the kinocilium with respect to the

total area of the cell biochemically and biophysically may affect attachment to the tectorial membrane. Thus cells that are signaling on the foreclaw display may grow large enough kinocilia to contact more strongly the membrane and vibrate with other cells responding. Thus the proximate mechanism is a density constant connection to other cells that respond which can give a larger reward sensation for the same behavior that is repeated in the coalitional group. Thus juvenile precocious play may be the beginning of climbing of the pleasure function which has inputs other than the foreclaw display. Thus as the turtle matures it can shift from an individualistic bias to teamwork in the climbing of the pleasure function.

chelonian vocalizations have been studied only occasionally (Campbell and Evans, 1967, 1972; Mrosovsky, 1972; for a review, see Gans and Maderson, 1973), and investigators considered tortoise vocalizations as simple noises involuntarily produced by copulatory movements (see Mrosovsky, 1972; Weaver, 1970;). Many studies (Campbell and Evans, 1967; Gulick and Zwick, 1966; Patterson, 1966; Ridgway et al., 1969; Wever and Vernon, 1956) ascertained that a number of tortoise species have a considerable auditory sensitivity to sounds below 1000 Hz, and this no doubt enables the animal to perceive many acoustic signals both on land and in water. Moreover, in a recent study on the terrestrial Testudo marginata (Sacchi et al., 2003), we showed that three features of male mounting calls (call rate, frequency, and duration) strongly correlated with male body condition and his mounting success (number of mounts achieved and number of females mounted). Behavioral Ecology (Jan./Feb. 2005) 16 (1): 301-308.

It occurred to me that the females gaped mouth may possibly be a sound communication back to male and this idea of sound communication occurred to Schnieder as well (with respect to vigalence behavior). Turtles do often release air under water. Is this accompanied with sound? So though turtles may not have language to create NBS binding contracts they may have complex abilities to distinguish different frequencies from different turtles. Turtles certainly do make sounds.

Here are two sounds recorded from Tracheyms on YouTube as well as what appears to be a similar sound under water from Chelodina. Another possible soud of the same type http://www.youtube.com/watch?v=p1UnlWpPY3M&feature=related

Youtube also a chirping baby Trachymes that seems to have a call that may also have its linguistic equivalents of a sound recorded also from Chelodina (needs a better visual presentation/figure to show possible comparability)

Could this open mouth below be a chirp at the end of series of bouts ( or has the turtle decided to eat instead)? http://www.youtube.com/watch?v=UvEjNkWo9Ms&feature=related

Some red-ears make a mewing sound either as young or perhaps when sick.

So population genetic roles (roughgarden Ackay PNAS 2011) that depend on matched foreclaw interactions might arise through sound modification of neural reactivity or possibly occur without sound vocalizations. The air sound of Trachymes contains frequencies I am fairly certain are produced by Chrysemys when biting at objects and simply ab initio

Wood turtles , known to associate in groups to hibernate,

have been recorded to vocalize while eating . Chelodina oblonga has a similar call as well.

Wood turtles also vibrate the ground with their front feet as birds do to attract worms to the surface. Could wood turtles have a means to have converted the foreclaw display (Including the position of the plastron) into a food gathering means they communicate to others aurally? Karen Davis has shown that turtles can learn feeding behavior from others so perhaps sound is the medium that turtles train others where and when to find food and form groups to hibernate? If this is the correct explanation then wood turtles could be investigated to dissect the difference between telenomic and telematic causes in the triple conintuum (two electrical directions and one mechanical) of the proposed proximate dynamics suggested here. Could the wood turtle stomp be the foreclaw display on land and a complex coalitional formation mechanism to share worms effecting communication about it? Is the Emydid foreclaw display ever associated with food? Like the social selection case with Trachemys, successful hibernation ability means longer survival and more likely-hood to be able to have more offspring successfully distributed in coming generations and thus if turtles attract others to groups through worm stomping then those groups social infrastructure could have been selected by stratigies to join the social group so doing. Thus it is not different mating stratigies of young turtles having to compete with old turtles but cooperation with older turtles to form coalitions that operates in turtle biology.

If sound behavioral ecology has evolved in various ways then it may be that the sounds associated with mating in Testudines may not be just for mating. There are cases where the these sounds are produced on shoes, onto turtles not in a mating position but in groups, single swimming sea turtle near the shore

, as well as deep underwater http://www.youtube.com/watch?v=DSEIfuonPTg&feature=related or above water

Ridgway et al 1969 green turtle 300-400Hz most sensitive useful span 60-1000Hz with a completely different sound (if I got this sampled correctly).

Female preference for fast-rate, high-pitched calls in Hermann's tortoises Testudo hermanni

Females prefer high-pitched calls and this correlates with smaller sized males. It may be possible that Testudo operates as was hypothesized with Trachymes with females behaviorally interacting with younger/smaller males but large males associating coalitions of long survivorship and more ability to leave offspring into future generations. Head bobing may be an effect on the vestibular system with inertial senses activated under territorial systems.Whether the NBS /NE active is a dyadic creating a polyad coalition or is polyadic in its raison de etra will depend on whether the pulsation is behaviorally different for triples than doubles in the continuum density and just not enough work has happened to say else still a runaway sexual selection processmay be operative. Ramming of the shell has been thought to be a trait necessary to imoblise the female but here one might consider that it causes afferent vestibular hair cell output into dopaminergic neurons which may efferently in pleasure affect inner ear hair cells priming them for auditory input to adapt the sound of the male creating a team of turtles that mate and listen to the stereotyped sounds of particular males. This would lead to social selection based on coordinated physiology of the vestibular and the inner ear hair cells. Indivdualism vs perfect team work would depend on the ability of a turtle ignore the imprinting that occurs during mating and could be heard 20 -30 meters around by other turtles. There is no need to have to hear two sounds simultaneously but simply the ability to revert to threat points which ignore the sound.

Sexual vs Social Selection in Tortoise Vocalizations associated with mounting. Galeotti et. al. 2005 have recently demonstrated for the first time that female Herman tortoises are sensitive to auditory playback and attracted by high-pitched and short duration calls of previously recorded mounting males. They cast their discussion of these results in terms of sexual selection but here, an alternative explanation based on a particular physiological hypothesis of sound energy transfer offers a social selection option to the conversation on natural and sexual selection in turtles. Although the ontology of social selection as a term can be confused because of different denotations (WestEberhard etc), Roughgarden ( 2012) has made clear how the term as applied in a two tier (behaviorevolution) process originally described in 2006 (Science) can be theoretically derived from perfect teamwork of mutual direct benefits. Here a suggested chelonian neurophysiology is offered to connote the meaning of the direct benefits and a practical hypothesis of social selection is presented for empirical testing. It is contrasted with the discussions on the difference of natural and sexual selection already undertaken for turtles. Here it is proposed that turtles have a common pleasure function in dopaminergic neuron topology and that the vestibular system can afferently send signals to this region which on pleasure can efferently effect differential sensitivity of the inner ear hair cells to auditory frequencies that are individually stereotyped. Perfect teamwork results when this connectivity that can thus be equilibrated in behavioral time is maintained during multiple repeated games. Threat points arise when individuals do not seek the pleasure gradient that maximizes both pleasures as one. This is not restricted to females hearing males but applies to other tortoises able to hear the calls in a region. There is a correlation between characteristic s of the call type and mounting success and tortoises geneally only vocalize during mounting intimacy. These facts lead G et al to suggest that vocalizations convey information about male features that females could choose qualitiatively. These choices are suggested to be individualistically benifical and not of a mutual direct benefit to both sexes even though there is no suggestion that males simply fight for females who are the prize. They suggest that the calls are costly condition dependent displays enabling males to succedd in sexual competition. In stead I suggest a cooperative teamwork functionality for the calls and suggest that biting the legs and butting the shell are not aggressive actions but positive reward sensations (through vestibular output) of mutual regard contributing the the attainment of perfect teamwork amonst individuals. This particular neurophysiological instatiation would preclude the tit for tat repeated dickering as suggested by Binmore in criticism of social selection . The results of G et al s work led them to say This experimental study showed for the first time in chelonians that female Hermann's tortoises pay attention to the quantity and quality of male mounting call, favoring fast-rate, high-pitched calls. On the other hand, males appear to detect conspecific calls, but they do not react to them in a consistent way, showing mixed responses to playback, with some individuals approaching and some retreating from the call of other males. Taken together, these findings weaken the hypothesis that male mounting calls may

serve an intrasexual function (i.e., avoiding rival interferences during mounting) and rather suggest that they are mainly directed to females, accomplishing an intersexual function toward the mounted female and possibly toward other females nearby. This is a general narrative of sexual selection to which social selection has been offered as a substitute (Por Royal Society 2012) Instead here I suggest that inner ear hair cells adaptation to individually stereotyped calls enables teams of turtles to form that increase the number of offspring in the next generation. Task allocation is not of females being choosy by running from males who have to chase them down but rather is a complex vestibular input into the brain that combines topographic impacts on turtle locomotion with physical impacts from another individual. It implies that male male fights may not competitive contests for females but rather vestibular cooperative behavior. The cost of the sound signal is irrelevant to amount of energy expended to acquire the region where the sound will eminate from. Thus the sexual selection view is not viable cast in terms of costly signaling and the dichotomy of sexual natural selection (Gibbons Lovich) also would not be operative since it would be teamwork rather than large amounts of individual threats that determine the variety of vocalizations behaviors (sounds) throughout the tortoises and thus predation pressure may not be the only other pressure than sexual selection ongoing in the terrestrial groups. The results would be to test if vestibular output effects input to inner ears If biting and shell ramming produces vestibular output Comparisons of predationpressure to others If females are choosing male quality through sound parameters.

African Spurred

chaco

red-foot

hindge-back

Egyptian These may represent different mechanisms of self-policing that prevents reversion to indivudalisitcally based threat points possible by zeroing out to self-motion perception via the vestibular system. Spite may arise by self-motion determined responses. It may be that reversion to individualistic behavior threat points may be associated with about 1,000HZ

Adaptation of vestibular signals for self-motion perception 1. 2. 3. Rebecca J. St George , 2 Brian L. Day and 1 Richard C. Fitzpatrick
1

+ Author Affiliations 1.
1

Neuroscience Research Australia and University of New South Wales, Sydney, NSW, 2031 Australia 2Sobell Department of Motor Neuroscience and Movement Disorders, UCL Institute of Neurology, Queen Square, London, UK Corresponding author R. St George: Neuroscience Research Australia, Barker Street, Randwick, NSW 2031, Australia. Email: r.stgeorge@neura.edu.au

1.

Next Section Non-technical summary The semicircular canals of the inner ear provide the brain with a sense of self-rotation and orientation. How they maintain a stable sense over time is not understood, as the signals from the canals have no fixed reference for rotation speed. We show here by stimulating the vestibular nerves electrically and by real rotation that a central adaptive process automatically zeros the vestibular signal to define the signal that represents zero rotation. In doing this, we also show that electrical stimulation generates a virtual signal of angular acceleration about an axis through the head. Knowing how these stimuli work and how the brain interprets them will assist in developing unique methods to investigate and manage a range of pathological conditions that produce abnormal sensations of movement and balance, and provide the understanding to develop novel virtual-reality techniques. Head bobbing, shell raming and biting may cause deviations from v estibular zeroing and if out put is to the pleasure center like neurons these othterwise catagorzied agonistic behaviors may be pleasurable casued by moving the otoliths relative to rest.

Experiments on waking curarized turtles showed that auditory representation is located in the mediodorsal zone of the tegmentum, in the torus semicircularis, which contains monomodal auditory and bimodal somatoauditory neurons. The somatosensory system is represented more widely and overlaps in the medial zones with the auditory system. Its focus is located in the lateral zones of the dorsal tegmentum (n. intercollicularis), where monomodal somatic neurons were found. Predominance of contralateral somatic projections was discovered. Frequencythreshold curves, obtained by analysis of evoked potentials, were flattened Y-shaped. The range of frequencies received was 406000 Hz and the range of optimal frequencies 100400 Hz. Responses of midbrain auditory neurons could be divided into three principal types: phasic, tonic, and bursting. Neurons with a phasic type of response were characterized by tuning to one optimal frequency, whereas most neurons with responses of tonic type were equally sensitive to two or even three frequencies. I. M. Sechenov Institute of Evolutionary Physiology and Biochemistry, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 14, No. 3, pp. 260269, MayJune, 1982.
Neurophysiology

Volume 14, Number 3 (1982), 191-198, DOI: 10.1007/BF01065122

Electrophysiological characteristics of representation of auditory and somatosensory systems in the turtle midbrain A. S. Khachunts

Keywords:

commissural connections; rhombencephalic tegmentum; extraocular motor nuclei; nuclei of the fasciculus longitudinalis medialis; pretectal connections

Abstract
Cerebellar and vestibular projections were investigated in the turtle Pseudemys scripta elegans following injection of 35S-methionine into the cerebellar and vestibular nuclear complexes at various locations. Fibers arising from the cerebellar nuclei were traced via the cerebellar commissure to the contralateral vestibular nuclear complex (particularly the n. vestibularis inferior and n. vestibularis ventrolateralis) and caudal rhombencephalic tegmentum. Ascending projections crossing the midline in the ventral isthmomesencephalic tegmentum terminated in the contralateral red nucleus and nuclei of the fasciculus longitudinalis medialis (flm). Vestibular projections ascending mainly via the flm terminated in the nuclei of the flm, the nuclei of the posterior commissure, and particularly the extraocular motor nuclei. Vestibuloocular projections arising from the rostral vestibular nuclear complex were almost exclusively ipsilateral; those from the caudal vestibular nuclear complex were bilateral. Evidence for a topographic organization of the projections to the trochlear and oculomotor nuclei was also obtained. There were some vestibular projections to the contralateral rhombencephalic tegmentum and n. vestibularis inferior. Spinal projections coursing within the ipsilateral ventral descending tract and the ipsilateral fasciculus longitudinalis medialis were found to arise from both rostral and caudal vestibular regions. The caudal vestibular nuclear complex in addition gave rise to fibers descending in the contralateral fasciculus longitudinalis medialis. Evidence for the existence of labeled fibers crossing at spinal levels was also obtained. Vestibulospinal terminations appeared restricted to the ventral horn.
The Journal of Comparative Neurology Volume 242, Issue 1, pages 102121, 1 December 1985 The cerebellar and vestibular nuclear complexes in the turtle. I. Projections to mesencephalon, rhombencephalon, and spinal cord 1. H. Knzle

Snapping turtle making sound Snapping of red eared slider with other underneath http://www.youtube.com/watch?v=3ij133XUua8

Spectrographs of the Sounds of Leatherback Turtles

www.jstor.org/stable/10.2307/3890632 by N Mrosovsky - 1972 - Cited by 11 - Related articles

Davis2009

There may be true triads as third ends up biting the displayer and the displayee ends up pursuing the biter here at the Prospect Park Zoo. This video looks like a cooperative triad a bit sequentially but is there sequestering instead (Ghislen) in all cases of triples 3 turtles http://www.youtube.com/watch?v=6459jxkWeDo&feature=related

http://www.youtube.com/watch?feature=endscreen&NR=1&v=mUDnE6dNqwI

http://www.suzannefredericqseaweedslab.com/. This video clip shows the courtship behavior of red-eared sliders (Trachemys scripta elegans), common turtles in the bald cypress (Taxodium distichum) swamp ("Cypress Lake") on the campus of the

University of Louisiana at Lafayette. Courtship and mating activities take place underwater. In this instance, two sexually active males simultaneously swim toward the female, stretch out their front feet, and flutter or vibrate their long claws on the female's head and neck in a courtship dance Mayr, Molecular Biology and the Mathematics of Selection Kant set himself up in opposition to Newton much in the same way that Einstein allied himself with Faraday and Maxwell. But what does a philosophical position entail for physical manifestations in the biology of form-making? It has long been suggested (Provine) that the different mathematical apparatus employed by Fisher or Wright does not implicate material differences in an understanding of evolution itself. If one looks beyond the physics of phase transitions between states of matter, the options opened up by the modern field of molecular biology permits a transition to a new unification of the multiplicity that dogs the repulsion of creationism from the plural science of evolution and enables a substantially new format for investigating the nature of lineages. Despite Mayrs strongly opinionated personality and often overly strong authorization of particular viewpoints he was on the edge of creating this philosophical insertion in his dissection of the meaning of teleology in biology. He apparently spent many hours thinking about Kant while consciously deciding that he had biology all wrong. He admired Kant but it seems that he never got to the point of fitting his interpretation of teleology within Kants discussion of Linnaeus and Erasmus Darwin as related to physics in Kants final unpublished (during Kants time) work, first published in English in the 70s. Kant had created a thought process in the Opus Postumum that permits one to sense (within the context that Charles Darwin was operating in ) philosophically (metaphysically) that, the mathematical differences between Fisher and Wright leads to different densities of populational-model thinking made mechanically available for influence on evolutionary dynamics (under infinite mathematical division per moving force) which when embodied by choices presented in the new discipline of molecular biology (which is often at odds with whole organism biology) forces a change both away from creationism and towards a new volume of testable hypothesis space without impacting social evolution however worldly considered. Many people have commented in the vein of MacNeil of Cornell that Mayr was just making up words (teleonomy and teleomatic) to avoid God in the room or a divine foot in the door (Lewontin) but as we can read from applying Kants transition from metaphysics to physics it is possible construct a reciprocal work programme for theoretical and empirical mathematical biology (that remains just as rigorous as its physical science counterpart) that circularly constructs enumerations and classifications of the forces moving between and connecting teleonomic properties with teleomatic universalities. Joan Roughgardens Stanford Lab has been promoting for almost a dozen years a challenge to Darwins sexual selection and it appears that unification of Mayrs view on somatic programming within his analysis of teleology can be incorporated into particular cases studyable in programmatic ways subject to social

infrastructure selection under syntheses of differences in competitive and cooperative game theory applied to lines of descent ascending through lists of forces categorized within expandable demes. Fisher made the theoretical analogy (Genetical Theory of Selection) between blending and particulate inheritance as one intuitionally associated with the kinetic theory of gases as under elastic vs non elastic collisions. In this Fisher set up the use of mathematics relative to selection and evolution to one of many different ways that moving forces may be enumerated as propagation continues. It is unclear how much the the 3-D nature of quaternionic algebra affected Wrights thinking but his use of path analysis provided him with a line of thinking to relate math to biology that did not need to imply the particular (non centripetal /centrifugal forces) with the motion under evolutionary change in the perception of the aggregated populational whole as did Fisher. Haldane associated with particular mathematical tools which constrains the ability to leverage his insights too forcefully in the continuum that the population is cut out of randomly (without respect to mutation per say). Mayr conceded that todaypg59 Towards a new philosophy of biology, we cannot understand development on the basis of purely physical laws as they were known to Kant. But I think I can show that by using Mayrs notion of somatic programming within Roughgarden et al format for social selection it is possible to understand some developments physically in a way that can be reinterpreted from Kants position through that of modern molecular biology. The first thing to do is to show where teleology itself junctions within biology. Modern Intelligent design fails for the same reason that Mayr thought Kant flunked out for biology because the purposive nature of organisms applies to invented and directly planned constructions conceptually like applying evolution to manipulate ecosystems, creating DNA computers to survey biological actions, and creating a community able to travel to the stars but not to analyses of paternity and the lines of descent that connect past forms of life into a grand dendogram tree classified and are mathematically operative in studies of heritability in evolutionary theory. It is the case contra Gihslen (Bioeconomicsw) that one should think of natural natural selection artificial natural selection and sexual natural selection because the systematics of the forces (different from the system of the forces (Kant on Linneaus) that social selection divides for different somatic programs molecularly in the instantiated behavioral equilibrium teleonomically while infinite theoretically teleomatically can only appear in nature in finite orderings or shapes/colors/body English. This type of thinking is a new philosophy of biology made in the transition to biophysics and shows that the observed need for a new theoretical biology in the 60s and 70s is indeed possible. Logic and reality are still just two different birds.

So the general idea is that turtles utilize the foreclaw pulsation to establish teamwork based pleasure coalitions that possess perfect (teleomatic) construction (all have the same pleasure gradient that changes under different adaptive uses of sound production and reception) due to the interaction of the mechanical and electrical properties on the nerve cells that are connected to the pleasure regions (teleonomically driving centers) in the brain even with a limited limbic system. th is an exceptionally

specific proposal and thus may seem unlikely but it is a case where social selection can be discussed with out direct involvement of parental care and thus demonstrates a case not considered in the conversation between social and sexual selection so far.

Much remains to be learned about this rather curious behavior in turtles, especially in nature. It appears that turtles may help to adjudicate the different theoretical ideas concerning social evolution (natural vs social vs sexual selection) generally. Does Sexual Selection 2.0 applied to females work for turtles with the foreclaw pulsation or is there a definable objective function that reflects changes in payoff matrices as turtles cooperate behaviorally and are rewarded sensationally efferently affecting the cooperative behavioral game in evolutionary time? Only more research can tell if Gibbons opinion that sexual selection is able to explain the length of the foreclaw used in the display no matter to what sense it is directed is true or not.

Juvenile red eared foreclaw pulsations http://www.youtube.com/watch?v=d9e2iwM2qP8 http://www.youtube.com/watch?v=-q1-mM2ImBQ&feature=fvwp&NR=1 http://www.youtube.com/watch?v=RPWa66GJiUc&feature=related

adults http://www.youtube.com/watch?v=tL9U_Q7gQfE&feature=related (positioning) Red eared copulation http://www.youtube.com/watch?v=Yl9O9LNup9w&feature=related

Juvenile and adult red-eared http://www.youtube.com/watch?v=4Rkne6noy6s&feature=related

References (not in proper form and not all included)

Gordon M. Burghardt. The evolutionary origins of play revisited: lessons from turtles. Animal Play: Evolutionary, Comparative, and Ecological Perspectives By Marc Bekoff, John Alexander Byers A.C. Crawford and R. Fettiplace ( The Frequency Selectivity of Auditory Nerve Fibers and Hair Cells in the Cochlea of the Turtle September 1, 1980 The Journal of Physiology, 306, 79-125.) C. H. Ernst 1971 Observations of the painted turtle, Chrysemys picta. J. Herpetol. 5(3-4): 216-220. Davis, Karen Marie, "Sociality, Cognition and Social Learning in Turtles (Emydidae). " PhD diss., University of Tennessee, 2009. http://trace.tennessee.edu/utk_graddiss/583 J. J. ART, M. B. GOODMAN Annals of the New York Academy of Sciences Ionic Conductances and Hair Cell Tuning in the Turtle Cochlea Volume 781, New Directions in Vestibular Research pages 103122, June 1996 Structure and Growth of the Utricular Macula in the Inner Ear of the Slider Turtle Trachemys scripta Stig vall Severinsen, Jrgen Mrup Jrgensen,and Jens Randel NyengaardStig vall Severinsen R. Fettiplace, P. A. Fuchs, Mechanism of Hair Cell Tuning Annu. Rev. Physiol. 1999 61:809-834 Johns Hopkins University Cochlear mechanisms from a phylogenetic viewpoint Geoffrey A. Manley doi: 10.1073/pnas.97.22.11736 PNAS October 24, 2000 vol. 97 no. 22 11736-11743 Nesse R.M. Social Selection and the Origins of Culture 2009 Evolution, Culture and the Human Mind phildelphia Pa Lawrence Erlbaum associates

Cell. Author manuscript; available in PMC 2012 January 7. Published in final edited form as: Cell. 2011 January 7; 144(1): 1626. doi: 10.1016/j.cell.2010.11.056 PMCID: PMC3038581 NIHMSID: NIHMS260995
Epigenetics in alternative pre-mRNA splicing Reini F. Luco, Mariano Allo, Ignacio E. Schor, Alberto R. Kornblihtt, and Tom Misteli
1 2 2 2 1

Sound Producing Mechanisms in Recent Reptiles: Review and Comment 1. 2. CARL GANS and PAUL F. A. MADERSON

+ Author Affiliations 1. Amer. Zool. (1973) 13 (4): 1195-1203. doi: 10.1093/icb/13.4.1195

Juvenile painted foreclaw http://www.youtube.com/watch?v=UNIExML05K0 map turtles foreclaw http://www.youtube.com/watch?v=BK1lN3_YyeU

Sound can be represented as actually infinite numbers of different frieze patterns layered in 3d Euclidean space. The ear hears by probing with 3 actually infinite points (associated with the

Quaternionic axes as in the point at infinity in complex number plane) by using the belt-trick to untwist the sounds rotations. This ability can be modeled with pairs of fundamental series (one end indicating amplitude and the other frequency) and the reverse used to point at any given rotation in any given density (magnitude of transfinites in the soundscape). Ordertypes thus span any reflections between the translations (convergence of the series) and rotations (3 points at infinity slicing rotations between *w and w). Cantor had differentiated the point at infinity of complex analysis with his newly proposed series of actually infinite numbers. Here we show how the ears morphological physiology operates with a materiality that connects both the points and the numbers substaintially. The expanse of the infinitely small to the infinitely large (of potential infinity) is Metaphysically contained within these layers by Kant under various Starken

or intensities of differently classified Linnean and moving force classes. It remains a need in science to have a classification of moving forces so that the infinite points can be enumerated into the layers per actually transfinite sets of freize patterns. Biology has not helped matters by thinking that forms can only not operate where teleology may have been thought. Thus Kants infinite in the Opus Postuum is largerly about the 3 point infinites connected to any transfinite infinite and not to Cantors transfinite infinite itself. Wesizacker mistook and confused the application of Cantors infinite into quatum mechanics thus by mistaking Aristotles and Cantors infinite. The quantum mechanical application of the actual infinite does need a particular transfinite but it is not the transfinities themselves (Russell attempt etc) but the ordertypes that bind a particular physical structure to the Euclidian space in which the object (on the large scale exists). It is possible that the model for the ear may help to develop ideas about space and time (with respect to mass) but I leaver that for another thought. The quantum mechanical use of number-classes depends on complete relation of potential infinities not on sensory modality specific projections.

The presentation of frequency and amplitude (waveform) in terms of back to back fundamental series (reversed and rotatable) shows what Cantor means by indicating that the infinitely small if they were real (they appear not when biology and physics is combined through chemistry) are different than the becoming infinitely small. As the belt-trick twists around a fundamental series pair the 3 points at infinity are crossed bodily as the relation between the frequency and amplitude becomes both infinitely small (frequency) and infinitely large (amplitude). (written Sept 3 2012 Brad McFall). The general history that discusses Cantors rejection of the infinitely small as real is mistaken and recent attempts to show that they are indeed objects crosses the phenotype genotype difference without regard. Robinson did not understand the difference between phylogeny (trees) and genotype (path analysis -Wright) and thus combines both cause and correlation if used for anything other than math. This is a legacy of Russells mistaken physical application of series of infinites generally started but not set within Kants pure physiology between moving force classifications and classified Linnean systems. There are new ways to understand the parallels of geometry through sense not simply apperceived

that takes evolution into account. Poincare did not do this. We can. Russell simply only took one possibility not an infinite number of them. So while we did not need or want metaphysical control when coming up with some/any physical idea of the actually infinite applicable to matter in combination through chemistry of biology (Robinsons mistake on infintesmal) and physics (Weisaker mistake on QM) it is need when evolutionary kinematics are dynamically combined to produce individual steps through transfinites over evolutionary time (dfferent forms formed). The difference of a transient meaning and a transient purpose were not understood by modern biologists and Mayr made this impossible to accord with the Darwinian tradition. As we develop DNA computer interfaces to tissue, enginner the ecosystem with applied evolution and travel and life off earth this will be enabled. Then perhaps the metaphyiscs will reappear as more and more transfinites are liked to different real not logical use of the 3 points (not one) at infinity compassed.

What is the origin of that information (1-D symmetries in force impacting space creating sounds and pressures and waves) and the development of canals to separate double quaternions into single quaterions (lateral lines from lancelets to morphologically made torus quaterions. With recapituatlions.) . It is possible to discuss explanatory and descriptive teleonomy within social selection. Teleonomic description depends on just what 1-D symmetries are in the environments past and present of the teleonomy so being explained. These symmetries may arise as by products of the behavior computer as somatic program recapitulated, through teleomatic necessities, or due to otherwise adapted features. The evolutionary tier is not preprogrammed but historical and depends on many things other

than just the behavior computers churning while changing. This churning and changing is in this case followable as the entropy measure of the teleonomic program from juvenile to adult behavior as the different quaternionic monomial become determinant for the entropy as the belt-trick is algorthmitized out of various homogneous polynomial functions of differing degress. The proximate reward sensation is the efficient cause of the ultimate acquisition of the trait. The proximate causes (quantum mechanical 4pie rotations in chemical bonds) mirroring this ultimate social selection have yet to be identified but are likely to be parts of the mechancanical electrical transduction means in the hair cells. This trait may contain a double quaternion use where individual threat points are operative asymmetrically thus providing a purely computational interval to friably separate material teamworks regardless of the physical waveforms (depending on how pleasure is neurobiologically instaintiated in turtles). Social selection thus can provide a kind of dynamics that is not reducible to the optimization framework suggested for universal biological dynamics.An optimization framework of biological dynamical
systems Ryota Horie Transcendetals categorize the countably infinite recovered in the social selection evo tier but remain logically within a superset of the non-countably infintly divisible sound types. The difference

between Williams and Mayr use of Pittenridgh depend on how infinity is found in the analysis. This places anti-Gould heterochornic space inbetween any velocities (accelerations retardations) and thus make a heterogenous set of polynomials but restricted in dynamics they can record given past recapitulations. Teleonomy Apropos: What is a goal of turtle ear organization? Ernst Mayr has made what he considers a factually based case that Goulds attempt to explain the mechanistic relationship between ontogeny and phylogeny via heterchorony is incomplete. (1994) He describes somatic programs as recapitulations and challenges molecular biologists to find the ultimate why of his reasoning and explain how it is that some ontogenetic developments require somatic programs and others do not. These somatic programs were described as a behavior computer in 1961. Though some attempts to bring this idea forward have appeared (Digitial ontogeny/phylogeny; Measure of Teleonomic entropy) many comments have be negative (Hulswit, Ariew,Francis,Thompson). I will show how social selection sensu Roughgarden is (dynamically)kinematically poised to differentiate the goal-directed processes implicated by the somatic program as behavior computers in the evolutionary tier by unconfounding the proximate reward sensation molecular biology from the behavioral equilibrium determining specific social infrastructures selected as adapted features by a direct application/instatiation/manifestation of the information metaphor Mayr introduced regardless of the teleomatic forces that the attraction and repulsion of the chemical bonds used/involved

necessitate. Thompson thinks that Pittenridghs idea that biologists who said, The turtle came ashore and laid its eggs rather than the turtle came ashore to lay its eggs meant to differentiate a description of an organization from an explanation of an organization. And that Mayr fails to do this. Mayr does say that proximate causes are purposive to the extent that computer programs are but that natural selection never is. Thus it will be possible to say that proximate reward sensation of turtles is TO have the most number of offsring and is done by the goal of either dipsplaying to the horizontal or the anterior posterior canals but because the molecular chemical bonds that that operatie the program that does the goal direction may be many different kinds (4pie rotations in the dual electrical mechanical continuum)Plural the ultimate somatic program retained reacapitualtionally is not to be a particular adaptive feature even though particular adapted features do arise as particular chemical attractions and repulsions are historically accumulated. Thus Huxleys random small order variation that is converted by the operation of selection into directional large-order variation evolutionary change along lines adapted to the improved performance of biological functions in a given enviornmenht does exist in turtle organization. Because there can be different molecular materials that control the quaternionic monomial constants during the attainment of specifc goal directed ends these ends nevertheless never are causally responsible for the current operation which is socially selected nevertheless. Social selection through specific somatic programs can be understood to fully unconfound the physiological mechanism from the evolutionary change.

Hulswit wrote, Mayr fails to explain what he means by causal. I can only interpret him as follows: a program is an efficient cause, which in combination with other efficient causes (the internal and external disturbances), completely determines the end state of the process or behavior. If so, then Mayr owes us an explanation of how under different circumstances, and thus, given different sets of efficient causes, the same program may lead processes to the same general end state. The behavioral tier equilibriums generally do this. More importantly: how can programs be considered as efficient causes at all? Or put more concisely: is there a theory of efficient causation that meets with the idea of a program? That seems unlikely, for efficient causes are singular events or facts, while programs are not. The different ways that the belttrick can be employed topologically/algebraically and with what particular quantum mechanical rotations provide, a rich field for meeting this requirement is extant. The singular quantum mechanical rotations may be in different chemicals but the 1-D freize pattern rotations are general for the particular interpretation of sound-vestibular functionality suggested here. There is also a possibility of using transfinite numbers of the patterns in the continuum (electrical-mechanical) should a better understanding of the relation of the telenomy and teleomatics was known.