This synopsis is designed to provide a quick overview of the contents of the new text, PROCESSES IN ANIMAL VISION which itself includes a major work on the FUNDAMENTALS OF THE NEURON as found in all animals.
The work was designed to satisfy two needs;
In accomplishing the above tasks, many new and unexpected phenomenon and mechanisms were documented. It also provided explanations for many inconsistent assumptions and positions in the literature. The resulting work is voluminous, approximately 2400 pages (and hundreds of new figures) in manuscript form. It also includes hundreds of equations defining the performance of individual mechanisms within the visual system.
To aid the reader, this synopsis is arranged to highlight different aspects of the manuscript. It includes the following major sections:
This synopsis suffers from some repetition. It is being edited continually. The repetitive portions should disappear as more chapters of the work are released in final form.
PART A develops the common architecture of all animal vision and demonstrates the potential for tetrachromatic vision in a plurality of animals. The large terrestrial animals and most arthropods are limited to different trichromatic subsets of the tetrachromatic spectra by evolution.
PART B introduces a new theory of the photosensitivity of the visual system based on modern physics that replaces the stereo-isomeric theory in use for the last 50 years. The previous mechanistic theory depended upon a putative structural change in the molecule named Rhodopsin. No investigation has ever been able to demonstrate the applicability of this molecule to the spectral performance of the chromophores of vision. The new theory is based on quantum-mechanical phenomena and leads to an explicit description of the photoexcitation process as well to the precise calculation of the absorption spectra of the individual chromophores as well as the detailed broadband spectra of the complete visual system.
PART C introduces a new theory of the neural system based on electrolytics. It replaces the previous ionic theory in use for the last 50 years. The previous theory depended on the ionic permeability of the membrane wall of neurons and an independence principle related to the flow of positive ions through that membrane. Current data shows that the intrinsic cell membrnae is totally impervious to ions because of its hydrophobic core and the independence principle has never been confirmed.
PART D provides a more precise level of detail describing the performance of the visual system. It provides a new Chromaticity Diagram to replace the current diagrams which lacks both a theoretical foundation and geometric conformality. By defining four regions of luminous performance, the functions and limits of adaptation and color constancy are presented explicitly.
This work originated in the 1960's with the realization that rhodopsin, as then defined, did not meet the requirements for being a chromophore. It was particularly deficient in the structural characteristics required of a good chromophore. This was understandable in light of the limited tools available in analytical chemistry of that period. The basic assumption had been that the residues of a destructive process could be easily returned to their original state and that state was a simple chemical bond involving only two components in a single molecule. This could be done conceptually and hopefully in the laboratory. The problem was the characteristics of the residues were not actually known. It was assumed that one of the residues was the alcohol or aldehyde of Vitamin A. The other residue was assumed to be a protein and was given the name opsin. Valiant, but unsuccessful, efforts were made to define the nature of the molecule and achieve the formation of rhodopsin in the laboratory.
A new class of retinoids was defined by the author at that time, the Rhodonines. This class met the requirements of physical chemistry and photochemistry for a high performance chromophore. However, it was difficult to obtain acceptance of the Rhodonines as a replacement for Rhodopsin within the vision research community. It was obviously necessary to place the material in a larger context to obtain such acceptance.
It was clear from work reported in the 1960's that the Rhodonines were actually present in the liquid crystalline state when employed in the vision process. This state accounted for their remarkably high absorption coefficient as well as their unique spectral absorption characteristics. However, this state of matter was virtually unknown to the Vision Research community.
More recently, the author undertook to determine the functional description of the photoreceptor cell and its relation to the photoexcitation process. This process produced remarkable results. It led to the discovery of the active mechanism involved in signaling common to all neurons. The Activa is an active three terminal electrolytic semiconductor device. The Activa is the electrolytic analog of the solid state transistor. A basic utility Patent has been awarded by the US Patent & Trademark Office relating to this discovery.
To properly understand the functional operation of the neuron, it is necessary that the reader be familiar with the concept of a "hole." In brief, a hole is an absence of an electron in a crystalline lattice. A hole can be described as a positive charge that moves in the opposite direction to an electron under electrostatic forces. The hole has only appeared recently in the literature of the bio-electrolytic chemistry field; a situation similar to that in solid state semiconductor theory in the 1950's.
A major corollary associated with the discovery of the Activa was that the fundamental active mechanism within a neuron is analog. This mechanism cannot be detected or confirmed by morphological or cytological means. It may not receive the appropriate emphasis but the vast majority of the neurons in any animal are associated with the sensing and inter-neuron processing of signals. These neurons all operate in an analog signal environment. It is only the signal projection neurons that employ pulse techniques to minimize energy requirements. They generate the "action potentials" so well known in neuroscience by the same simple process used widely in solid state transistors.
Following the discovery of the basic signaling mechanism of the neuron, its location within the neuron but external to the nucleus became obvious. The general location was already described in the literature. Several discoveries occurred following the discovery of the basic signaling mechanism. The most important was the discovery that there were two primary applications of the Activa in vision, and in fact in all neural systems.
The Activa constitutes the signal manipulation device at the dendrite-axon interface inside a given neuron.
The Activa also constitutes the primary signal transmission device at the axon-dendrite interface between neurons, the so-called synapse.
These two discoveries place the signaling function of the neural system in animals on a firm electronic foundation. They also lead to a clear definition of the hybrid role of the Node of Ranvier as a synapse internal to a specific neuron.
To understand the operation of the photoreceptor cells in more detail, it is necessary to understand their role in a larger context. This requirement led to the development of a much larger block diagram, and subsequently detailed circuit diagram, of the entire visual system. In doing this, the obvious problem was the lack of consistent information about the visual system in the literature. Conflicts abound for at least two reasons; the tendency of individual investigators to adopt "floating models" that only apply to their specific area of interest, and the general use of only algebra based tools in the analyses. Even where more appropriate mathematics was employed, they were still inadequate. The visual system is a very sophisticated system. It uses many of the most sophisticated methodologies known to man at the start of the 21st Century. Failure to recognize these mechanisms and methodologies leads to an inadequate understanding of the overall process. The visual system employs a number of time related processes that have not previously been addressed in the literature. To understand these processes, it is necessary to employ "complex algebra" in the differential equations arena. Employing these techniques provides the complete solution to the overall photoexcitation/de-excitation process within the Outer Segment of the photoreceptor, including the variable time delay with excitation that has so confused earlier investigators. They also led to the understanding of the adaptation process and the unique electrical topology found within the photoreceptor cell.
The definition of the active mechanism within the neuron and the use of more sophisticated mathematical tools has introduced a new understanding of the "excitability of the neuron" and the crucial role of the Node of Ranvier. Rather than the outer membrane of an axon being an intrinsically excitable structure, it is the junction between two such membranes that introduces the phenomena known as "transistor action." Every neuron includes at least one active device exhibiting this phenomena, as does the structure known as the Node of Ranvier and all other axon-dendrite interfaces. The conclusion is drawn that the entire animal neural system is electrolytically based and active. The primary role of chemistry is in the electrostenolytic processes providing electrical power to this extensive electrical network.
The development of a detailed circuit diagram of the visual system has produced new insights as the architecture and signal handling techniques used in the neural system in general. The most important findings were in two areas; the definition of the specific signaling paths associated with both achromatic and chromatic vision and the mechanisms of perception used to interpret the signals passed over these paths. A related major finding was that there are two distinct entry points for visual signals into the cortex of the brain and the area known as the "primary visual cortex" is not the most important one in many respects.
With the detailed circuit diagram of the visual system available, it is possible to provide a new series of performance descriptors for the visual system (essentially transfer functions of the overall system) based on a much stronger theoretical foundation. These descriptors are generally compatible with previous descriptors but offer much more detailed insights into the underlying mechanisms. However, they are generally based on different theoretical principles than the conventional wisdom as summarized in the documented list of paradigm shifts between this work and that wisdom.
The ability of the electrolytic interpretation of the operation of the neural system, including the operation of the synapse, to describe the detailed performance of the system appears to leave the alternate chemical hypothesis untenable.
When reviewing the vision literature, one cannot overlook the plethora of inconsistent terminology used by the community. This situation is illustrated by the confusion of letter symbols used to describe elements of the retina and the mid-brain by various researchers. There is a similar plethora of inadequately defined, and frequently conflicting concepts used to define the visual process. The problem appears to be compounded by the part-time nature of the typical academic researcher who cannot commit significant blocks of his time exclusively to his research intersts. It has also been compounded by the historically poor preparation of the researchers in the field of mathematics. These conditions appear to have caused the development of a variety of incomplete theories that exhibit two major difficulties. They usually use inadequate mathematical tools and invariably are based on partial or "floating models." The most glaring example is the continued reliance within the community on the concept of linearity as a keystone of the visual process. These conditions have led to the development of a series of "Standards" by an international body that are based entirely on a limited conceptual foundation acquired through inadequate experimental activity. Although part of this inadequacy can be assigned to the natural advance of technology with time, it does not justify the failure of the community to update those standards during the last 50 years.
Although morphology and physiology have existed side by side for a very long time, it is apparent that morphology, and cytology, have reached their zenith and are now holding back the field of vision. The functional aspects of vision are found in the realm of electro-physiology. Contrary to the beliefs of some individuals, form follows function in vision as in nearly every field.
The lack of a lead or central researcher (or a series of such researchers over a period of time) with a global view of the visual system has also contributed to the current state of knowledge in the field. This state continues to exhibit local "schools" with a strong research interest in only a narrow sector of the visual process but an inadequate perview of their interest in terms of the the overall context of vision.
A new comprehensive theory of the visual process is presented. It can best be described as an ORTHOGONAL ZONE THEORY based on the terminology of previous theories. The extensions involve both greater interior detail as well as the addition of additional zones to the earlier zone theories.
The goal has been to present an overall view of the visual system in a defendable mathematical context and a global scientific framework. This goal has required the introduction of techniques and mechanisms not normally found in the literature of vision. This has been particularly true in two areas, the definition and detailing of the initial photodetection process and a similar detailing of the mechanisms of neural signal transmission. In both cases, the dominance of chemically based concepts is shown to have impeded progress. The description of the visual system, including the neural system, as an entirely electronic, more precisely electrolytic, based system leads to much greater insight into the operation of the visual system than any chemically based theory can offer.
A major finding has been that color vision is pervasive throughout the animal kingdom, has been since the earliest evolutionary times, and is frequently tetrachromatic. The performance of the human eye is atypical with respect to its spectral capability, along with other large land mammals, due to other adaptations introduced to satisfy environmental needs.
A second finding removes the widely held view that the photodetection mechanism differs significantly between the invertebrates and the vertebrates. By developing a complete phylogenic tree, it is shown that there are at least three (not two) fundamental types of eyes. It is also seen that all three use the same functional architecture (topology) in their photoreceptor cells. This functional architecture is obscured by the great variation in morphology of these cells.
The ORTHOGONAL ZONE THEORY is able to properly place the trichromatic, Hering and other zone theories in a proper (and larger)context. It is also able to establish the features and shortcomings of the C.I.E, the Munsell, the Swedish natural, and the OSA color systems based on theoretical grounds. These same theoretical grounds lead to the presentation of new luminosity functions, a new chromaticity diagram, and a more comprehensive adaptation characteristic, covering both dark and light adaptation. Finally, it provides a bridge between past studies of the eye and the brain as separate individual entities in the visual system.
In the development of the ORTHOGONAL ZONE THEORY of vision, an entirely new FUNCTIONAL THEORY OF THE NEURAL SYSTEM evolved. It is based totally on the electrolytic nature of the neuron and the electronic nature of all signals within the system. This theory explains the operation of the neural system of all animals down to a level of detail that any current chemically based theory can only aspire to achieve. As an example, the action potential of the ganglion cell is shown to consist of two separate waveforms in the temporal domain. The first is a rising exponential waveform exhibiting one time constant and the second is a falling exponential waveform exhibiting a second time constant. The transition is abrupt between these and it does not occur at the centroid of the combined waveform. The action potential of the Node of Ranvier is seen to be formed using a similar but distinctly different methodology.
A major demonstration of the FUNCTIONAL THEORY OF THE NEURAL SYSTEM is that the synapse, along with the Node of Ranvier and an internal junction only hinted at in the literature are totally electronic with respect to their signaling properties. The concept of a neurotransmitter is not required.
A number of other cherished but archaic concepts of vision are also discarded in the development of these works. Probably the most venerable is that of a dichotomy between types of photoreceptors, the rods and cones. The theory demonstrates in excruciating detail that there is only one functional type of photoreceptor cell and that it is associated with one of four types of chromophore. These chromophores are sensitive in the ultraviolet, the short, the medium and the long wavelength portions of the visual spectrum of light.
PART A opens with a major discussion, Chapters 1 and 2, devoted to the physical environment and the phylogenic tree of the animal kingdom. It is necessary to explore many of the ramifications of how animals exist in their environment in order to provide a framework for a global analysis of the visual process. An initial finding was that the common division of animals into two major branches, protostomic and deuterostomic, is not an appropriate one based on vision. It became clear that vision began with a common ancestor, probably planaria, and has evolved along three different paths in conjunction with the three large animal classes; Arthropoda, Mollusca and Chordata, more colloquially described as the insects, the molluscs and the vertebrates. This common ancestry does not support the common reference in the literature to a basically different operational mode for the photoreceptors of Arthropoda compared to Chordata.
The amount of differentiation within each class has caused significant areas of overlap in the visual processes of these classes. However, the class relationship must be respected if a true understanding of the visual process in a given species is to be obtained.
Before proceeding, a series of simple experiments are provided to help the reader evaluate a group of commonly held misconceptions about the human visual system. The goal being to establish a common framework for further discussion. This area also includes a review of many of the laboratory procedures used in the past to investigate the visual process. A number of suggestions are made to improve the experimental design of many investigators in order to insure a wider and more general applicability of their results.
The material also addresses a series of generic subsystems applicable to the visual system and shows how these systems play different roles in different species.
This presentation also demonstrates that first order or so-called Gaussian optics can not be relied upon in vision research. Gaussian optics are limited to the paraxial condition, i.e., less than one degree from the optical axis. The fovea is slightly more than five degrees from the optical axis. This fact and the high curvature of the retina calls for the use of a full optical analysis, including fifth order aberrations, in visual research.
Chapter 3 discusses the retinas found in various visual systems. It stresses their commonality where appropriate, and also highlights their differences as a tool to further understanding. It also introduces the extremely important subject of the metabolic and hydraulic aspects of the operational retinal. Later, these subjects are shown to be of critical importance to the understanding of the overall visual process. The outline of the overall visual architecture begins to appear in this discussion.
Chapter 4 focusses on the photoreceptor cell of vision. To maintain a manageable discussion, it proceeds to focus on the photoreceptor cell of chordate vision. An initial effort is spent on terminology. The problem of terminology is endemic to the subject of visual science. This is because of the many disciplines involved and the tendency of individual investigators to introduce "local concepts" to help interpret their findings. Later, it will also be seen to be due to the critical lack of a precise definition of even the basic term color.
The Chapter drills down through the anatomy, morphology, cytology and molecular chemistry of the photoreceptor cell in considerable detail. It also stresses the importance of the support system provided to the photoreceptor cells by the retinal laminate and the biochemical matrices found within the individual laminates. Later, this support system is shown to be critically important to the overall visual process. The discussion develops the dominant characteristic of any neural cell, its electronic, or electrolytic nature. With this electronic nature clearly in mind, the actual electrolytic topology and topography of the cell are developed. This results in a description of the unique electrical topology associated with the sensory neurons of the animal nervous system. It also hints at the very sophisticated nature of the adaptation amplifier found in the human visual system. The actual performance of this circuit is discussed more fully in PART E
This Chapter concludes with a detailed examination of the life cycle of the photoreceptor cells and the retinoid material employed in the chromophoric portions of these cells. It is here that the fact that the Outer Segment of the photoreceptor cell is located external to the actual cell first becomes important.
PART A concludes in this Chapter with a schematic of the interface between the photoreceptor cells and the RPE (retinal pigment epithelium) that summarizes both the static and dynamic aspects of the photoexcitation process.
Part B focuses on the photochemistry of the visual process in detail. Chapter 5 develops the fundamental chemistry and the quantum physics of the chromophores of vision. The first four major sections are devoted to terminology and scientific background. They address the specific quantum physical characteristics of retinoids when present in the liquid crystalline state. It is shown that there are a unique set of parameters required of any good chromophore. It is also shown that the structure known as rhodopsin does not exhibit these properties but that these conditions are found in a sub-family of the retinoids, the Rhodonines, that are directly derivable from retinol, Vitamin A. The theoretical spectrums of the Rhodonine family of retinoids are shown to match the best measured spectrums of animal vision in all four spectral regions, the ultraviolet through the red.
The distinction between the isotropic absorption spectrum (peak at 493 nm.) due to the molecular absorption dominant in dilute solutions and the anisotropic absorption (peak at one of four different wavelengths) due to resonant absorption by retinoids in the liquid crystalline state is stressed. The anisotropic nature of the resonant absorption spectrum is quite apparent when using the suction pipette technique to measure the in-vivo spectrum of an Outer Segment. The spectrum of all photoreceptors exhibits a peak wavelength at 493 nm when exposed to irradiation transverse to the axis of the Outer Segment. When exposed to radiation parallel to their axis, the same Outer Segments exhibit their biophysically relevant spectral absorption characteristic. A discussion is also provided explaining how past laboratory extraction methods, which have been unsuccessful in isolating the chromophores of vision and demonstrating the spectral characteristics of the visual chromophores, can be modified to isolate and confirm the presence of the actual chromophores of vision. The method is based on recrystallization hile in the liquid crystalline state.
PART C follows as an unanticipated drop-in part. It concentrates on the newly discovered functional mechanism within the neurons of all animals with particular focus on the visual system. To appreciate this PART, Chapter 8 begins with an expansion and a codification of the terminology of the neuro-scientific literature. It is shown that the fundamental operational structure of the neural system is not the neuron but an active electrolytic semiconductor device accompanied by its input and output structures. Cytologically, this structure consists of an electrolytic input conduit (usually described as a dendrite), an electrolytic output conduit (usually described as an axon) and the active electrolytic semiconductor device between these two conduits. The latter structure is unknown in the current literature and has been named an Activa. This fundamental structure is most clearly seen along a projection neuron. It is typified by splitting the so-called interaxons of the axon in two. The fundamental neural structure is then seen to consist of the Activa located at a Node of Ranvier, the adjacent (dedrite like) portion of the distal interneuron, and the adjacent (axon like) portion of the proximal neuron. Every synapse in the system can be similarly represented as the junction between two conduits connected by an Activa. A similar structure is found between every dendrite and axon of a neuron.
Based on the above finding, the so-called neuron cell is actually a metabolic biological structure designed to support one or more (complete or partial) fundamental operational structures of the neural system.. The neuron is not a good morphological descriptor of the operational units of the neural system. The typical projection neuron usually contains as many as 100 more fundamental operational units. The Chapter includes a discussion of a "fundamental neuron" that can be described as a single fundamental operational structure supported by a single soma. The Chapter also develops the electrical parameters associated with such a fundamental neuron and how it is supplied electrically from an electrostenolytic process involving the glutamate family of biochemicals.
Chapter 9 proceeds to develop the cytology and the electrical topology of the more complex elements of the neural system, especially the photoreceptor cells and the projection cells (typically called ganglion cells in the peripheral neural system. The photoreceptor cell is shown to incorporate unique circuit features shared only among the sensory neurons of the body, as opposed to the signal manipulation and projection type neurons. This unique structure accommodates the input signals accepted from the actual sensory structure bonded to but not part of the photoreceptor cell, but known historically as the Outer Segment. It also provides the unique variable gain amplifier that is the heart of the adaptation process. It is this process that generates the so-called adaptation phenomena, light adaptation and color constancy. The Chapter includes a brief codification of the parameters of the active devices, the Activas, found in the neural system. These tabulations are similar to those for man made transistors, the immediate cousins of the biological equivalent Activas.
Chapter 10 re-examines the morphology of the neuron based on the improved understanding of the electrophysiology of such units developed above. It develops the detailed morphology of the various types of neurons based on their electrical topology and topography as well as their cytology. The unique morphology of the lateral cells (the horizontal and amercine cells) is explained based on their unique performance requirements. They are shown to exhibit two fundamentally different input structures that allow the internal Activas to act as both inverting and non-inverting amplifiers simultaneously, thereby providing an electrical differencing circuit.
The synapse, the Node of Ranvier, and the internal junctions within a "fundamental neuron" are shown to all be functionally identical. They are all electrolytic (electronic) interconnections containing an Activa.
This chapter provides a complete electrical circuit diagram for a fundamental (straight through) visual sensing system, from the Outer Segment of the photoreceptor cell to the stellate cells of the cortex. This basic diagram is expanded to include a variety of other electrical signal paths associated with the visual system. The symbology used is drawn from the conventional symbols used by electrical engineers. However, make no mistake, these symbols are representing biologically identifiable mechanisms. In the case of the individual Activas, there is no need to replace them with an equivalent circuit containing a great number of electrical elements. The Activa is a fundamental electrolytic circuit element.
A key expansion of the above electrical circuit diagram is to provide the complete circuit diagram of the servomechanism at the heart of the Precision Optical System (including elements of the auxiliary optical system).
The electrophysiology of the neuron is explored in detail. The applicability and limitations of the voltage clamp technique are also discussed in depth. It is shown that the precise use of this technique requires considerable more attention to detail than found in the current documentation. The detailed schematic of the voltage clamp technique illustrates the five major variants in the laboratory technique that explain the frequent inconsistency in the data from peer investigators.
One of the major findings of this work concerns the "ion pump." The finding involves the separation of the electrical functions of the fundamental neuron into two related but distinct processess. The first has to do with the biasing of the active devices within the fundamental neuron. The second has to do with the response of the membrane wall to changes in polarization across it. The finding is that ions are not transported across a neural membrane by a chemical process to create an electrical bias and sustain the operation of the neuron. On the contrary, only electrons are transported across the membrane in response to an electrostenolytic process on the surface of the membrane. These electrons provide the long term polarization of the electrolyte (plasma) internal to a given neurite or axon. It is the polarization of these conduits that bias the Activa at each junction into its operating range. In response to the operation of the Activa, the electrical bias of the axonal electrolyte may be changed on a short term basis. This short term change requires the transport of charge across the membrane wall in order to sustain electrical equilibrium. Because of this charge transfer, the number of ions on each side of the membrane is changed. Thus, it is the change in the potential of the axonal plasma relative to the exterior plasma that causes the (apparent) transfer of heavy molecular ions across the membrane. The actual physical transport is limited to electrons. No actual ions transverse the membrane. It is the electrical potential across the membrane that is the source of the (apparent) ion pump, and not the ion pump that is the source of the potential.
A second finding is that the axolemma of a neuron, a simple biological bilayer of known electrical properties, is not an active or "excitable structure." It is the junction between the dendrolemma and the axolemma of the neuron that produces "transistor action" through well understood quantum mechanical mechanisms that creates the "excitable" phenomena associated with the neuron. This fact accounts for the difficulty of repeating experiments of the voltage clamp type unless great care is taken to remove all of the dendrite related structure from the sample neuron. This procedure deactivates the active device. The subsequent tests record the response of a passive structure to pulsed electrical excitation in accordance with the laws of passive electrical networks.
The resulting topography and topology of the fundamental neuron presents a much more detailed perspective of the operation of the neuron. This has many impacts on the laboratory test environment. It is shown that the so-called Voltage Clamp Technique of the literature has not been adequately documented in the past. By following the documentation in the literature, it is possible to create at least five different voltage clamp test configurations. This has been the source of a great deal of difficulty in the laboratory in the past and hampered the independent confirmation of the early experiments.
The impact of pharmacological preparations on the neural system are seen to be primarily associated with the electrostenolytic processess on the surface of the neurons responsible for the electrical potentials of the various electrolytes of the neuron. By interfering with these processess, the operation of the fundamental electrical circuits can be impacted or stopped.
Part D takes the knowledge developed in the earlier parts to develop a detailed understanding of the signaling environment of the visual system and closes with an interpretation of the resulting psychophysiology applicable to all animals. A special emphasis is placed on the human because of the breadth of the literature available in this area. The performance of the system is explored one stage at a time and then one circuit at a time. Chapter 11 is devoted to a discussion of the concept of modeling of biological phenomena. It is followed by Chapter 12 focusing on the Signal Detection process, Chapter 13 on the Signal Manipulation process within the retina, and Chapter 14 on Signal Transmission (Projection)to the brain. Chapter 15 concludes with an important review of the signal interpretation functions of the brain, both in the old brain and the cortex.
Chapter 11 provides a discussion of the importance of the tradeoff between using a large signal model, which always applies, and using a medium or small signal model that only apply under restricted conditions. Chapter 11 also provides an in-depth review of the literature related to the Electroretinograph (ERG) in order to develop a coherent presentation on the subject and rationalize much of the literature with the actual mechanisms underlying the measured data. The fundamental waveforms defined in this work are compared to the recorded waveforms of the literature. This methodology provides a more fundamental understanding of the measured waveforms.
A series of simplified block diagrams of the visual system are presented. They can be used profitably when exploring particular aspects of vision.
Chapter 12 is extensive and detailed. Because of its great importance in vision, the complete electrical circuit of the photoreceptor cell is presented in detail. This circuit is presented as an overlay to or adjacent to the cytological drawing of the cell in order to aid understanding. The physical separation between the Outer Segment and the rest of the cell is highlighted from a number of perspectives. The critical electrical separation of the Outer Segment from the rest of the cell by the Outer Limiting Membrane is illustrated.
Other than the beauty associated with the overall circuitry of the cell, the features of the adaptation amplifier within the cell are unique and noteworthy.
Chapter 15 concentrates on Higher Level Processing, particularly those steps leading to, and including, the Perceptual processPART E presents both global and specific analyses describing the visual system in animals. This analyses also includes Sections on failures in the system as well as abnormal or aberrant operation of the visual system. It closes with a discussion of areas of further study and a preliminary summary of productive experiments.
Chapter 16 begins the PART by establishing a mathematical framework for assembling the material in the previous chapters into a cohesive whole. The result is a series of comprehensive equations, based on a single end-to-end model, describing the visual process in animals. These equations can be simplified by holding a variety of different parameters constant. The resulting simpler equations provide precise descriptions of a wide variety of individual phenomena found in the experimental literature. Several sections of the Chapter address specific phenomenum.
The section on adaptation describes the complete adaptation process, both light and dark. It provides the first known theoretically based description of the phenomenum of dark adaptation. The important revelation is that this characteristic is not the result of two exponential processes due to two conceptually independent mechanisms, typically described as rods and cones, that originated with Hecht in the 1930's. It is shown that the phenomenum is due to a single process described by a second order differential equation. The mathematical solution of this equation describing the process of dark adaptation is the product of an exponential and a sinusoidal term. Such an equation is frequently described as an exponentially damped sinusoid. However, because of the relative amplitudes involved, it is more appropriate to describe the function as a sinewave modulated exponential. This function fits the available data on dark adaptation precisely, including the rise in (or at least leveling of) the response in the vicinity of 25 minutes after the cessation of illumination found in the data of Hecht and others.
Chapter 17 is designed to illustrate a new set of functional descriptors of vision, and specifically human vision. The chapter presents a completely new set of theoretical luminous intensity related descriptors along with the procedure for degrading them to obtain facsimiles of the current C.I.E. Standards. Similarly, it presents a completely new set of theoretical chrominance related descriptors along with a procedure for quantifying the Munsell Color System to absolute spectral wavelength. This material provides an absolute definition of the terms "color" and "white," and an assortment of spectrally unique names for individual colors.
The chapter also develops the fundamental characteristics of a three dimensional SENSATION SPACE. This three-dimensional sensation space is not compatible with a spherical coordinate system.
The chapter also provides a series of descriptors of the transient performance of the visual system. Specifically, it provides the first known description of the complete adaptation cycle, from dark to light and return. Both the light and dark adaptation phases are shown to agree with the empirical literature. However, the underlying mechanism of the dark adaptation phase is shown to be fundamentally different than that suggested in the literature.
Chapter 18defines a wide variety of idiosyncracies and malfunctions of the visual system, with particular emphasis on those associated with color vision. Many of the indiosyncracies are seen to be due to compromises made in the design of (optimization of) the the signal encoding and signal projection techniques used in vision. The abnormalities of color vision are seen to depend on a great many variables. Many observed syndromes may be due to multiple underlying mechanisms. The variety of these failure modes illustrates the difficulty in relying on the early literature attempting to explain inferences between these failures and specific abnormalities in a specific samole of genetic code.Chapter 19 provides a review of alternative theories of vision. It shows that both the Young-Helmholtz and and Hering theories are subsets of the ORTHOGONAL ZONE THEORY presented here. In the neural area, the excited membrane theory is found inadequate when compared to the "transistor-action" based junction theory.
The work concludes with:
The scope of the complete ORTHOGONAL ZONE THEORY is great. It encompasses a great many aspects of vision. It includes a range of static relationships as well as a range of dynamic relationships. The latter involve temporal intervals of microseconds to hours and involve a great breadth of technologies. This range includes many functions designed to maintain the homeostasis of the system; the circulation and the transport of critical constituents, metabolism, and the renewal process within the retina. The latter process extends from disk formation through phagocytosis.
The following material will present a few key vignettes defining individual features of the visual process as interpreted by this ORTHGONAL ZONE THEORY.
With the above foundation, it became possible to understand the operation of the photodetection process. This explained the operation of the Outer Segment associated with the photoreceptor cell as well as the operation of the photoreceptor cell itself. The results were very satisfying. The results led directly to the Photoexcitation/De-excitation equation that described the generator waveforms found at the terminals of the photoreceptor cell/Outer Segment association. The word association is used because it became clear that the Outer Segment is not an integral part of the photoreceptor cell. It is a structure extruded by the photoreceptor cell and, therefore, found outside the cell membrane.
With the modern tools of physical chemistry and electrical engineering available, it was possible to define the structure of the Outer Segment in considerable detail. This led to the determination that the spectral absorption characteristics of the chromophores, when deposited as a liquid crystal, on to a substrate in more detail. This solved one of the classic problems of experimental electrophysiology as it applies to vision. The Outer Segments exhibit a distinctly anisotropic absorption spectrum. The distinction involves two separate mechanisms. Each of the chromophores in the Rhodonine family exhibit a common isotropic absorption spectrum with a peak absorption wavelength near 500 nm. This spectrum is due to the monopolar characteristics of the molecules conjugated hydrocarbon backbone. Simultaneously, when in the configuration found in the Outer Segment, i.e., a liquid crystalline state of flat surfaces, the molecule exhibits a second highly directional absorption spectra particular to its unique bipolar physical chemistry and conjugated hydrocarbon backbone. This absorption spectra has a strong peak at its characteristic chromophore wavelength to light applied axially to the Outer Segment. Thus the dichotomy between the results of reflective spectrophotometry through the iris of the eye and the results of transverse spectrophotometry performed in a variety of electrophysiological experiments.
With an understanding of the quantum mechanical operation of the chromophores applied to the substrate of the disks, it became possible to explore the interface between the Outer Segment and the photoreceptors in more detail. The ramifications of this exploration were significant. They included:
the confirmation of the unique two-exciton process found in the long wavelength photoreceptor channel of animal vision and predicted from the laboratory results in man regarding the photopic and scotopic spectrums as a function of illumination level.
the demonstration of the fact that there are no achromatic photoreceptors in animal vision.
the recognition of the extremely high effective quantum efficiency of the visual process in animals--rivaling successfully the highest quantum efficiency achieved in man-made semiconductor based sensors (>90-95%) and far higher than that achieved with photoelectric tubes (including the photo-multiplier types).
the location of the adaptation amplifier that; controls the signal level in the remainder of the visual signal processing chain and accounts for the remarkable dynamic illumination range of the animal eye. [The term dynamic is used to indicate that the instantaneous range is limited to only about 120:1 within a total range exceeding 1,000,000:1.]
the corroboration and explanation for the ability of the animal eye to detect individual photons. This corroboration also defined explicitly the fact that the chromophores of vision are not energy detectors--they are quantum detectors. The use of equal energy spectral signals in vision research is not appropriate and leads to fundamental errors in calibration and measurement if not accounted for.
the recognition of the second class of Activa within the photoreceptor cell which acts as a distribution amplifier serving the synapses associated with the pedicle.
With the nature of the signals available at the pedicle of the individual photoreceptor cells understood, it was possible to categorize the photodetection channels of vision and to explore the signal processing functions of the retina more precisely. This has led to a clear distinction between the three broad classes of signal processing within the retinas of all animals, with emphasis varying between species, the luminance, chrominance and appearance channels. It is not possible to explore the appearance channel of animal vision extensively at this time because of the limited scope of the literature. However, it was possible to explore the luminance and chrominance channels in considerable detail. This effort uncovered four extremely important features of all animal vision:
the fundamentally tetrachromatic aspect of animal vision, the ultraviolet capability being lost in atmospheric animals employing complex eyes due to the absorption characteristics of the cornea-lens combination.
the generation of both the exact photopic and exact scotopic spectral response (as well as the intermediate mesopic and the hyperopic spectral responses) of the complex eye based on the input of signals from only the chromatic signal channels associated with photodetection. There was no need for a so-called rod sensor, an achromatic sensory channel.
the fact that the typical animal eye incorporates three chrominance difference channels, only two of them being active in atmospheric animals with complex eyes--such as humans.
the fact that the signal projection elements of the visual system were fundamentally transparent to the signals being carried. This resulted in a quasi-perceptual plane located at the input to the ganglion cells that closely mimicked the putative perceptual plane within the brain. The failures in the signal projection elements to be transparent are the source of most of the erroneous perception related flicker phenomena observed in human vision.
With a quasi-perceptual plane recognized within the retina, it becomes possible to explore the signal projection elements of the visual system in more detail. This exploration quickly defined the nominal role of the ganglion cells, the mode of signal encoding used and certain secondary characteristics of interest.
The ganglion cells are seen to be simple reflex oscillators of a well known type. They can operate in the mono-pulse, bistable or astable mode depending on the function required, and occasionally on how they are treated (abused) by an electrophysiological experimenter. The ganglion cells associated with the chrominance channels, believed to be the midget ganglion cells, normally operate in the astable (or free running) mode. The ganglion cells associated with the chrominance channels normally oscillate at a pulse to pulse interval of about 33 milliseconds, near the fusion frequency of the human eye.
The ganglion cells associated with the luminance channel, believed to be the parasol type ganglion cells, operate in a somewhat more complex mode. In the absence of photoexcitation, they operate in the astable mode. Upon photoexcitation at a level exceeding the ganglion cell threshold, they generate a single pulse, known as an "action potential." If the photoexcitation remains above the threshold or increases, the ganglion cell generates additional pulses with a time interval between pulses indicative of the strength of the signal.
The signal in the luminance channel is seen to be encoded using time delay between pulses as the "carrier." This signal encoding mode is known as time delay modulation, a common form of phase modulation. It is important to note that the modulation is not frequency modulation. Frequency modulation cannot carry information in as compact a form as phase modulation. Normally frequency modulation cannot carry any information concerning the first pulse. The initial pulse in the luminance channel is used in conjunction with the tremor to indicate alarm or change. The same type of encoding is used in the chrominance channels except for an offset. The time duration before or after the occurrence of the nominal pulse is used to encode a bimodal signal. When encoding high saturation color information, the resulting signal is asymmetrical. This asymmetry is the source of most distortions associated with flicker phenomena.
It is very easy to implement a signal decoding circuit in the brain using a single Activa per projection path. Subsequent signal processing in the immediate vicinity of the decoding Activa is analog in nature. Only when information must be transmitted over distances longer than a millimeter can one be assured that an additional encoding step will be involved using a projection neuron similar to a ganglion cell.
The significance of the mid-brain to the overall visual process has been widely overlooked. Although its significance has been explored recently with regard to the LGN and the presence of signal paths to and from the pretectum have been documented for a long time, the literature still provides an elementary view of the importance of this area. The fact that it plays a crucial role in the manipulation of time dispersed signals is virtually unreported in the literature. Similarly, the fact that it plays a crucial role in the dynamics of the line of fixation of the visual system is also underappreciated. These shortcomings are unfortunate since time dispersal is the key to understanding the role of the midbrain in vision.
Time dispersal, introduced in the topography of the axons of the ganglion cells of the retina and removed by Meyer's Loops between the LGN and area 17 of the cortex, is a key tool in the design of the visual system.
Time dispersal also plays a key role in the Precision Optical System of vision. This system incorporates what has long been known as the Auxilairy optical system (AOS) of the mid-brain into the primary servomechanism loop used to control the involuntary aspects of the line of fixation of the eyes. By relying on time dispersal, the signal processing within the servo loop can be performed without requiring significant and explicit short term storage of signals within the AOS. The AOS plays several other critical roles with respect to the line of fixation since it receives inertial stabilization signals from the vestibulary system and voluntary pointing instructions from area 7A of the cortex.
Time dispersal plays a key role in the time-dispersal encoding technique employed throughout the signal projection systems of the neural system of animals. The visual system takes full advantage of the time dispersed signals in the LGN in order to extract the signals needed to perform stereopsis and avoid parallax errors in the areas of object space of primary interest to the animal. To a large extent, these tasks are performed withn the laminated functional structure of the LGN. Signals received at incrementally different times are compared to ascertain the displacement and velocity of small elements of the visual fields shared by the two eyes. The resultant signals can be used to obtain both the pointing instructions for the individual eyes that will minimize parallax errors within the fovea and the rate of motion instructions required by the AOS to establish the proper angular rates for the eyes via the Precision Optical System servomechanism loop.
Until recently, the measurement of visual signals within the brain was extremely limited due to limitations on experimentation with humans and on the limited ability of man to communicate with animal subjects. Recent major advances in radiometric techniques such as the MRI are providing tremendously valuable new information relative to the time relationships between various visual signals. One of the most exciting, and in a sense most perplexing to previous theorists has been the fact that many significant signals appear in the higher cortical centers of the brain before they appear in area 17, the "primary visual cortex." Although the pulvinar pathway between the pretectum and area 5 of the cortex has been known for over 50 years based on morphology, its significance in vision was not appreciated. It is now clear that one of the functions of the Precision Pointing System and the AOS is to extract signals from the foveola of each eye and transmit the key features of these signals directly to the feature extraction engine of the cortex located within area 5. Because of the limited spatial extent of the signals from the foveola, these signals are not mapped spatially within area 5. They are treated as features in vector from their initial arrival. Other signals extracted from the LGN and transmitted to the area 17 include significant spatial information. As a result, this information is mapped spatially on the surface of area V1, a portion of area 17 based on a different nomenclature. As the spatial information is converted to vector form, within the cortex, the need for spatial mapping disappears. This fact is recognized by the lower and lower degree of spatial correlation relative to object space found in areas V2, V3, V4, etc.
The entry of important visual information into the cortex by a path other than via the area 17 undermines the current literature attempting to define two parallel processing paths beginning in the LGN and continueing via area 17 to the frontal lobe of the cortex. The cortex (at least of higher primates, which technically does not include the rhesus monkey, Cercopithecoidea macacus) is seen to be organized according to a star architecture to minimize signal delay between related feature extraction engines and command generation engines. This architecture is compatible with other sensory inputs than visual and provides an overall minimum time delay between signal input, cognition, and response command generation.
The star architecture of the cortex is an asynchronous architecture that allows different engines of the cortex to use different amounts of time to complete their tasks without compromising the ability of the animal to respond quickly when necessary. It accounts for the fact that the animal may take evasive action before it can identify internally, or verballize externally, the character of the threat.
There are several major theoretical results:
the demonstration of the critical role played by the liquid-crystalline state of matter, and the unique electrolytic properties associated thereto, in the vision and neural systems
the demonstration of a new photochemistry for the photodetection process in all animal vision
the discovery that the neural system consists of large array of electrolytic conduits separated by active electrolytic amplifiers defined as Activas and supported metabolically by the soma and nuclei of cells.
the discovery of the Activa as the primary mechanism of the neural system in all animals
the recognition that the biological bilayer membranes used to form the conduits of the neural system are not permeable to ions and that the diffusion laws of Nernst, Donnan, Goldman and Hodgkin & Huxley do not apply to them.
the discovery and demonstration of the fact that the visual and neural systems do not employ dissipative thermodynamic principles.
the demonstration of the fact that feedback, in its external form, is not used in the visual process
the recognition that feedback, in its internal form, is key to the operation of the photoreceptor cells
The discovery that the entire neural system operates under reversible thermodynamic principles in order to achieve its unique level of thermal efficiency is fundamental. This discovery also places the subject of bio-energetic fuels to support the neural system in an entirely different light. It provides conclusive evidence that the fuels represented by the glutamates, GABA and the non-protein amino acids, such as glycine, do not participate in the signal detection, processing or transmission process in vision but are critical to the power supply function associated with the electronic nature of the neural system. In this role, the density and consumption of the bio-energetics associated with a photoreceptor cell is proportional to the illumination level. The density and consumption associated with other visual cells is less direct. These bio-energetics are required by every neuron. They are known to be ubiquitous throughout the body.
All of the above and following postulates are overwhelmingly supported by the available literature and the experimental work of many individual investigators.
Based on the above finding, described in detail and documented in this work, it is possible to develop several important practical results and applications. The overriding result of this work is a contiguous theoretical model applicable to the Visual Process in All Animals with its associated equations and demonstrations of applicability.
The most important practical result is the elimination of the putative achromatic "rod" from further research interest. All animal photoreceptors employ one of four chromophores, three in atmospheric complex eyes, from which all achromatic and chromatic information is extracted. The second most important result is the recognition that vision does not involve linear signal processing. The common linearity laws do not apply. In fact, a majority of the visual signal path employs "large signals" at all times. This makes even the imposition of linearity based on the small signal model inappropriate at light levels above that equivalent to the full moon. Analysis of the visual system at the research level requires the use of logarithmic algebra. The third fundamental result is that external feedback is not employed nor is any form of retro-signaling, such as dendrite to axon or dendrite to dendrite.
Although it may continue to be useful for engineering purposes to use empirical values for the spectral performance of the animal eye, a single equation is now available that defines the radiometric performance of the eye under all slowly changing irradiation conditions. These conditions range from the hyperopic thru the photopic and mesopic to the scotopic. A perturbation analysis of this equation at a given irradiance provides clear indications of the rigor required in laboratory investigations if useful data is to be obtained. A simple calculation also becomes available to indicated the error inherent in using a fixed spectrum photometer, particularly one employing an energy sensitive detector, to simulate actual visual performance.
A single pair of equations is now available that defines the chromatic performance of the human eye. With the actual equation available describing the spectral absorption performance of the eye, it is no longer adequate to assume the trichromatic generalization is true. This generalization is based on a group of linearity laws that are not applicable to the vision process when studied from the research perspective. Specifically, what is known as The Color Equation, the linear sum of the contribution of each chromophoric channel as a representation of the total chromatic experience, is now archaic. At the research level, logarithmic algebra is required.
Imposition of logarithmic algebra has many benefits. It negates the need for such empirical tools as "tristimulus values, including putative real and imaginary spectrums. The word real in the previous sentence refers to "realizable but synthetic spectrums" as used by the C.I.E., not to the actual spectrums of the chromophores of vision. The use of logarithmic algebra also obsoletes the C.I.E. Chromaticity Diagram for research purposes. A New Chromaticity Diagram suitable for research purposes is now available which involves orthogonal coordinates in either two or three dimensions. Only the two dimensional version is needed for humans and other complex eyes within the atmosphere.
The New Chromaticity Diagram solves the century old battle between the Hering and Young-Helmholtz schools. The ORTHOGONAL ZONE THEORY provides the theoretical foundation upon which to evaluate the proposals of both schools and the associated shortcomings of each theory. This foundation shows the idiosyncracies of the fundamental axes that has led to the debate over these two theories.
The theory defines and The New Chromaticity Diagram illustrates that the sensation of "White" is uniquely defined in perceptual space. The white of perceptual space is transformed uniquely into object space when the object space luminance represents an equal photon flux spectrum. If the large field object space luminance is not represented by an equal photon flux spectrum, the adaptation amplifiers in each photodetection channel will attempt to compensate for this fact. This compensation manifests itself in two ways:
The C.I.E. has defined a variety of specific photon flux spectrums using the term luminants, i.e., luminants A, B, C and the more recent D series. The D series is flexible. It uses a subscript to indicate the color temperature of the source in hundreds of degrees Kelvin. Whereas the D65 luminant is most frequently mentioned in the literature, it is the D70 illuminant that most closely represents the equal photon flux condition. It is this condition that the generic visual system was apparently optimized for.
The Theory, model and equations clearly demonstrate that the linear addition of pigments and lights in object space must be clearly differentiated from the chromatic response of the organism in perceptual space. These two representations of color are only related by a very complex mathematical transform at the research level. The transform can be simplified for engineering applications as long as the appropriate caveats are noted.
The theory and equations also define explicitly how the sensation of color is lost as the luminance level is reduced, leading to achromatic perception at scotopic illumination levels. As long as the adaptation amplifiers are operating at less than maximum gain, they produce essentially saturated chromatic signals at the input to the chromatic differencing circuits. The result is a variation in perceived hue that can be accompanied by high saturation levels. At lower illumination levels, signals at the input to the chromatic differencing circuits are reduced in amplitude. The resulting difference signals are also reduced in amplitude. As a result the signals applied to the midget ganglion cells are reduced and the level of modulation of the encoded information is less.
The type of modulation used does not convey information about hue and saturation separately. The system was implemented on the assumption of constant maximum saturation level under normal (photopic) conditions.When detected within the brain under less than photopic conditions, only reduced levels of saturation can be perceived. This eventually leads to a perceptual range that is limited to white and black. This occurs at scotopic levels. No achromatic photodetectors, such as "rods," are required in this process.
The signal processing related to the geometry of scenes has not been treated in as much detail as the photometry and colorimetry of the visual process. This is due to a paucity of detailed information to draw on. This paucity is in turn due to the complex relationship between the spatial and the temporal performance of the visual system. The block diagrams and schematics applicable to the appearance signal processing channels have been outlined. These outlines should encourage further experimental activity.
Individual circuit diagrams are presented for each stage of the visual process, photodetection, or signal detection; luminance, chrominance and appearance signal processing; and signal transmission or projection. These actual biological circuit diagrams can be emulated using electrolytic (either biological or non-biological) semiconductor or solid state semiconductor devices. These circuits were emulated in the solid state for the military during the 1970's. The purpose was to demonstrate feasibility of a 100,000,000 detector mosaic for Inter-continental Ballistic Missile (ICBM) tracking.
With a detailed model of the visual system of any animal available, it is possible to explain in detail most of the many transient and unusual effects recorded in the literature under the names of various investigators. These proper name references can now be dropped in favor of more specific mechanism related names.
The overall model and the equations provide a new and unique ability to aid the clinician in the isolation of medical problems. The model also provides for the first time a definitive description of the types, sources and potential medical cures for a wide gamut of color vision abnormalities. The model and equations are able to help immensely in the attack on the more intractable problems of current visual pathology.
The fundamental premise that the neurological system is electrolytic in nature imposes a requirement on the oRTHOGONAL ZONE THEORY to offer an explanation of the basic action of pharmacological agents on the neural system. Such an explanation is readily available. It involves the electrostenolytic processes occuring at multiple locations on the surface of every neuron. These processes provide electrical power to the neural system via the glutamate cycle of metabolism. Any interference with the operation of this cycle will have the effect of impeding the transmission of neural signals. This is most easily done by impeding the flow of the necessary metabolites to or from the surface of the cells or by scavanging the constituents involved in the process and impeding the appropriate chemical reactions.
This work has provided a new and much broader understanding of the Process of Animal Vision. However, to appreciate the visual process, certain revisions in the conventional wisdom must be made. These revisions involve our understanding of certain fundamental physical and mathematical concepts, certain realizations about the basic processes of vision, and many of the tools used in visual research. These revisions have led to a variety of important discoveries.
The ability of this work to mathematically define a more precise set of visual performance functions for the human eye, and show how these functions can be degraded to produce the current C.I.E. standard functions, is very strong support for the accuracy of the signal generation, signal manipulation and signal topography proposed in this work.
The ability of this work to mathematically generate an accurate synthetic ERG, and define where each feature in the waveform originates (to within microns) is strong vindication of the entire electrolytic hypothesis of this work..
The support for the original definition, developed by the author in the 1960's, of the actual chromophores of the visual process provided by this cumulative work is of great satisfaction.
The discovery of the active electrolytic semiconductor device supporting the entire neurological system of biology is an even greater, and unexpected, reward for the last six years of intense devotion to this project. A United States Patent has been awarded recognizing and protecting this discovery.
To understand the operation of the animal visual system, it is mandatory that some broadening of the accepted concepts be employed:
+It must be recognized that there is a fourth state of matter, the liquid crystalline state, that is present ubiquitously in living organisms and exhibits many unique properties in the operation of the organism.
+ It must be recognized that Gaussian statistics are not appropriate to many visual processes and events. Gaussian statistics are an approximation, for large aggregate quantities, to the more fundamental Poisson statistics. Poisson statistics hold for small quantities as well as large in the absence of other "control parameters." Statistical events involving quantum events include an additional control parameter, Pauli's Exclusion Principle. These events, such as photodetection, are represented by Fermi-Dirac statistics. Statistical events involving biological growth also involve a (different) control parameter and are represented by Poisson statistics in logarithmic quantity space. The resulting statistics are frequently approximated by log-normal statistics (or log-Gaussian statistics). The data can be approximated by a "bell shaped curve" on a logarithmic horizontal axis.
+ It must be recognized that the thermodynamics taught in general education, and involving a Carnot Cycle, is not the only form of thermodynamics. The Carnot Cycle is based on the assumption that some heat is generated in the process under discussion. Many reversible electro-chemical reactions do not release heat. These reactions operate in accordance with the laws of reversible thermodynamics. The electrostenolytic processes employed in the neurological system involve reversible thermodynamics.
+ It must be recognized that the presence of a material ubiquitously in the neurological system does not mean it is a participant in the signaling function of neurons. In the neurological case, there are two environments in the vicinity of the neuron interconnections (synapses and Nodes of Ranvier). In the perinodal area, the dominant material is a minute liquid crystal of pure water (hydronium) and the signal carrying charges are electrons and holes. In the paranodal area, the most important materials with respect to vision are the bio-energetics participating in an electrostenolytic process and related to the glutamates and butyrates (simple amino acid derivatives). The charge carriers in the electrostenolytic processes are electrons and ions. The transport of ionic materia within the paranodal region does not constitute signal transmission.
+ It must be recognized that vision, and the operation of the neural system in general, is a non-linear process involving logarithmic mathematics.
+ It must be recognized that many families of Retinoids, and the properties of the liquid crystals, were unknown at the time that the basic assumption was promulgated that the photoreceptors of vision involved Vitamin A in simple molecular union with a protein. Definition of the Rhodonine family of Retinoids provides the actual chromophores of all vision when present in the liquid crystalline state and in electronic contact with the neurological system (or other discharging mechanism).
Based on the above facts, it is possible to summarize the fundamental findings of this work.
Linearity is not a valid concept related to the visual process. A caveat applies only to very specific experiments that are totally restricted to small signal conditions. An example would be small signal frequency response measurements made under static illumination conditions.
Additivity as a concept derived from Linearity, is not applicable to the visual process.
Superposition, based primarily on Linearity, is not applicable to the visual process, even under small signal conditions when investigating color perception.
The photodetection process involves a mixture of linear and square law processes.
The high degree of negative internal feedback within the photoreceptor cells allows the entire remainder of the visual system to operate in a mode defined by a fixed amplitude range, which must be considered highly non-linear.
It must be recognized that the tools developed in the 1930-60 time frame by the C.I.E. do not rest on the foundation and the realizations outlined above. Whereas the C.I.E. tabulations and diagrams are adequate for normal engineering applications, they are not adequate for research purposes.
Based on this work, the various empirically based and discrete luminosity functions, photopic and scotopic, can be replaced by a theoretically based and continuous luminosity function that can be considered to include four discrete regions; hyperopic, photopic, mesopic and scotopic.
Based on this work, the empirical C.I.E. Chromaticity Diagrams can be replaced by a theoretical New Chromaticity Diagram for Research. This New Chromaticity Diagram is free of the many ambiguities and imprecise aspects of the old diagrams.
Based on this work, the old empirical "dark adaptation curve" can be replaced by a new theoretically based "full adaptation function" that incorporates both the old "dark adaptation curve" and the virtually non-existent "light adaptation curve" as special cases. The resulting function dispenses with the notion of "rods" and "cones" and correctly describes the transient response of the eye under a variety of conditions involving illumination intensity and spatial position. The function clearly presents the state variable nature of the visual process.
1. The discovery of the actual chemicals (the Rhodonines) involved in the chromophores of vision and their presence in the liquid crystalline state of matter. This discovery replaces the less defined description of the "rhodopsin concept" in vision.
2.The development of the inherent tetrachromatic visual capability of all phyla of animals. The human being among the unfortunate few who are only able to use a trichromatic portion of this capability.
3. The discovery of the active electrolytic semiconductor device found in all neurons and crucial to their operation.
4. The discovery that the neurological system in animals employs mechanisms based on reversible thermodynamic principles. The mechanisms do not employ a Carnot Cycle, do not dissipate heat, and do not conflict with the Second Law of Reversible Thermodynamics.
5. The discovery that the Node of Ranvier is the prototypical synapse. It is the simplest and easiest to study of all synapses. Functional synapses, by one name or another occur between neurons as well as within cell bodies.
The book contains 19 generously illustrated chapters, includes a six-page summary of the tabular parameters of the human eye, an extensive glossary and approximately 10 appendices. A more complete description of the book, a summary of the hypotheses within the Theory, and sample chapters are available at the Vision Concepts website.
James T. Fulton1 (949) 759-0630 (Pacific Time)
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