Spacenoology

Here is temporarily posted the text of the thesis from the site: http://www.dissercat.com/content/sravnitelnaya-morfologiya-i-opticheskie-svoistva-glaz-bryukhonogikh-mollyuskov-stylommatopho
Year:
2007
Author:
Shepeleva, Irina Pavlovna
Academic degree of:
Ph.D.
Place of defense of the thesis:

Moscow
Discipline Code WAC:
03.00.08, 03.00.13
Specialty:
Zoology
Number of Pages:
147
Contents:

Introduction.

Chapter 1. A comparative essay on the structure and optics of the eye of invertebrates with emphasis on chamber eye gastropods.

1. Types of invertebrate eyes.

1.1. Simple eye.

1.1.1. The light-sensitive eyes.

1.1.2. Pigment glasses.

1.1.3. Pinhole eyes.

1.1.4. Camera eyes.

1.1.4.1. Aquatic animals.

1.1.4.2. Terrestrial animals.

1.1.5. Camera eyes with reflective layer.

1.2. Evolution of simple eyes.

1.3. Compound eyes.

1.3.1. Apposition eyes.

1.3.2. Superposition eyes.

1.4. The evolution of compound eyes.

2. Camera eyes gastropods.

2.1. General structure of the eye.

2.2. Characterization of cellular and acellular components of the eye.

2.2.1. Tentacular epidermis.

2.2.2. Perioptichesky sinus.

2.2.3. Eye capsule.

2.2.4. Cornea.

2.2.5. Lens.

2.2.6. Vitreous.

2.2.7. Retina.

2.2.8. Additional OPC body.

3. The spatial resolution and sensitivity of the optical chamber eye.

3.1. General Provisions.

3.2. Factors affecting the spatial resolution.

3.2.1. The number of photons absorbed.

3.2.2. Spherical aberration.

3.2.3. Chromatic aberration.

3.2.4. Diffraction.

3.2.5. The structure of the retina.
Introduction:

Approximately 570 million years ago in the history of the animal world was one of the most spectacular events in just a few million years - a moment in geological standards - appeared on the Earth most of the animals, known to this day. Animal Ecology - their physical habitat and their interaction with each other to implement various forms of behavior - led to the evolution of orientation, which resulted in the emergence of a wide range of eye types of structure. In invertebrates they account for no less than nine (Warrant, 2003).

Type Mollusca - monophyletic line of invertebrate animals (Cook, 2001), which has successfully penetrated into many habitats of oceans, seas, freshwater and terrestrial (Pasternak, 1988). A variety of devices, eye clams ranges from the most simple pigment cup with saucer (Patella sp.) To the chamber of the eye lenses, the lenses look like fish, octopus in (Octopus sp.) (Land, 1981), ay bivalve genus Pectunculus Area and there are even the most complex eyes that resemble insect apposition eyes (Land, 1984).

The richest by the number of species (over 30 thousand) class type is the class of Mollusca snails - Gastropoda. Originally Gastropoda - sea animals, but in the course of phylogenetic development of several groups of mollusks of the class adapted to terrestrial life. Thus, the gastropods were the only group of mollusks that have occupied the land (Fechter, Falkner, 1990). Traditionally, according to the classification of messenger (Messenger, 1981), mollusks Class Gastropoda identify three types of eye: 1) the pigment cup (marine limpet Patella sp.); 2) pigmented glass, filled with gel material (abalone Haliotis sp.); 3) eye chamber (marine mollusk perednezhaberny Strombus sp.). Dominant in these molluscs are chambered eyes with the cornea and lens, separated from the retinal layer of the vitreous body.

Some researchers believe that the snails define objects through chemical and feeling that the eye chamber can provide only the reaction of phototaxis (Eakin, Brandenburger, 1975; Audesirk, Audesirk, 1985; Emery, 1992; Chase, 2001). However, the implementation of this form svetozavisimogo behavior can be achieved by not fotosensornoy ocular system (Vakolyuk, Zhukov, 2000; Millott, 1957, 1967; Steven, 1963; Stoll, 1975; van Duivenboden, 1982) and thus does not require specialized optical devices and relatively developed retinal eye chamber (Zhukov, Baikov, 2001). Meanwhile, in behavioral experiments on the ability of gastropods to the discrimination of visual stimuli shows that some of these animals can distinguish between the geometric shapes that resemble objects of their natural habitat. Marine molluscs perednezhabernye Littorina punctata (Evans, 1961), L. irrorata, Tectarius muricatus, Turbo castanea (Hamilton, 1977; Hamilton, Winter, 1982, 1984), freshwater pulmonary Planorbarius corneus (Zhukov et al, 2002), Lymnaea stagnalis (Andrew, Savage, 2000) and ground lung Achatina fulica (Zhukov , Baikov, 2001), Otala lactea (Hermann, 1968), Helix aspersa (Hamilton, Winter, 1984) using visual information for orientation in the environment.

As seen on the visual abilities of gastropods formed fairly conflicting views. The reason is that until now we have a few scattered data on the visual system of these animals. Analysis of the literature showed that the number of works devoted to studying the possibilities of the visual system of the gastropods are very small and contrasts with the large number of studies carried out in arthropods and vertebrates. Judging by the publications of interest to the visual system of gastropods originated in the late XIX century. The greatest number of experiments to the period from 1965 to 1979. In the 80s and 90s there is a tangible decline in the number of published papers, and from 2000 to 2006. appeared in print a few articles. At the moment, for the vast majority of the investigated gastropods only data on total and fine structure of the eye (Henchman, 1897; Prince, 1955; Clark, 1963; Tonosaki, 1967; Stensaas et al., 1969; Hughes, 1970; Jacklet et al. , 1972; Mayes, Hermans, 1973; Eakin, Brandenburger, 1975; Kataoka, 1975, 1977; Zunke, 1979; Eakin et al., 1980; Katagiri, 1984, 1986; Tamamaki, Kawai, 1983; Tamamaki, 1989; Katagiri et al., 1995; Blumer, 1998, etc.). Simultaneous study of the morphology and optics of the eye chamber devoted only a few works: L. littorea (Newell, 1965; Seyer, 1992), Agriolimax reticulatus (Newell, Newell, 1968), Strombus luhuanus (Gillary, Gillary, 1979), Pterotrachea sp. (Land, 1981), L irrorata (Hamilton et al., 1983), S. raninus (Seyer, 1994), Ampularia sp. (Seyer et al., 1998), L. stagnalis, PI. corneus, Radix peregra, Physa fontinalis, Trichia hispida, Cepaea nemoralis (Bobkova et al., 2004 (I); Gal et al., 2004 (II)). Observations on the relevant forms of behavior in mollusks, for which data are already available on the structure and optics of the eye, even more scarce and confined L. irrorata (Hamilton, 1977; Hamilton, Winter, 1982), L. stagnalis (Andrew, Savage, 2000) and PI. corneus (Zhukov et al, 2002) Thus, it is clear that in our knowledge of the visual system of the gastropods, there are many gaps, and the question of its functional significance is still open. Meanwhile, an evolutionary approach to the study of visual function requires a corresponding detailed studies of this rather large group of animals that occupy different habitats, showing a variety of forms of behavior and have, accordingly, different sensory organs are arranged (Zhukov, 1990). A comparative study of the visual system is ecologically isolated species of gastropods may show the influence of environmental factors on its development in the phylogeny and to identify the functional significance of this system for animals. Such studies will determine the possible range of adaptive changes of the sensory system of gastropods (Hamilton, Winter, 1984). In addition, information about the device and the optical eye will provide a basis for appropriate behavioral experiments, the results of which, in the end, and build our judgments about the possibilities of the visual system of gastropods.

Purpose - a comparative study of the morphology and optical properties of the eye of terrestrial gastropods, different light regime of habitats - Arion rufus, Perforatella incarnata, Helicigona lapicida and have similar lighting preferences - Arianta arbustorum, Cepaea hortensis.

Experimental study of the problem:

1. To study the general morphology of the eye.

2. Describe the dioptric apparatus of the eye: to describe the components that form the image, and estimate the focal length.

3. Characterize the light-sensitive eyes and svetoizoliruyuschy apparatus.

4. Rate spatial resolution and sensitivity of the optical eye.

5. Establish the extent to which structural and optical properties of the eye and visual function to determine the adaptive devices for visually impaired at the level of sensory organs.

6. Identify key areas of change in eye device and the properties of their components due to the transfer of molluscs in the phylogenetic development of the water in the terrestrial environment.

Scientific novelty of the work.

1. First studied the morphology and optical properties of the eye of terrestrial gastropods, Arion rufus, Perforatella incarnata, Helicigona lapicida, Arianta arbustorum and Cepaea hortensis.

2. For the first time the comparative analysis of structural and optical properties of the eye with a variety of gastropods, and similar lighting preferences.

3. In terrestrial gastropods, first demonstrated the dominant role in the lens focusing light.

4. In gastropods, first discovered four morphologically distinct types of photoreceptor cells in mikrovillyarnyh retina J. lapicida: one type of photoreceptor of the first type and three types of photoreceptors, the second type. It is suggested that such a variety of cell types can provide color vision shellfish.

5. For the first time shows the relationship between the brightness of the preferred habitat of mollusks and optical sensitivity of the photoreceptors of the first type.

6. For the first time tracked the main directions of restructuring of the eye due to the transfer of molluscs in the phylogenetic development of the water in the terrestrial environment.

Theoretical and practical importance of work. Obtained by the methods of light and electron microscopy data and the results of making a significant contribution to the study of structural and optical properties of the peripheral visual system of mollusks Class Gastropoda. This information is important for knowledge of the functional significance of the visual system for this group of animals. The results of significantly extend the existing ideas about the possible range of adaptive changes in the peripheral visual system of gastropods to terrestrial habitats and to habitats with different light conditions. The results provide a basis for ethological studies examined in this study of mollusks, and allow us to determine directions for further research of the visual system of gastropods. The data presented are necessary for understanding the evolution of the visual system and can be used for Malacology, evolutionary and comparative morphology and physiology.

The provisions that are brought to a defense.

1. In all studied species of molluscs are the dominant lenses optical components. Lenses A rufus, N. lapicida, A. arbustorum and C. hortensis, obviously, have a gradient of refractive index. The lens P. incarnata, apparently, is optically homogeneous.

2. In A. rufus, P. incarnata and N. lapicida, with different lighting preferences, found an inverse relationship between the brightness of the preferred habitat, and optical sensitivity of the photoreceptors of the first type. In A. arbustorum and C. hortensis, living under the same lighting conditions, found similar values ​​of the optical sensitivity of the photoreceptors of the first type. In all studied species of molluscs between sensitivity to light photoreceptors and brightness of the second type of habitat is no connection has been observed. Within the eyes of each type of shellfish seen the following relations: a high optical sensitivity of the photoreceptors of the first type / low optical sensitivity of the photoreceptors and the second type of low spatial resolution of the photoreceptor of the first type / high spatial resolution of the photoreceptor of the second type (with the exception of the second type of photoreceptor of the second kind in N. lapicida) . It may be more sensitive photoreceptors of the first type operate at low light levels, and the photoreceptors of the second type - with a bright light.

3. Major changes in the structure of the eye caused by the passage of mollusks in the phylogenetic development of the water in the terrestrial environment, can be traced in the direction of thickening of the cornea, the acquisition of a soft consistency and ellipsoid lenses, as well as reducing the distance between the lens and the retina to the minimum possible. Apparently, the ancestors of perednezhabernyh clams have inherited the dominant role of the lens in focusing the light and the gradient of refractive index of the lenses.

4. In the formation of the eyes of mollusks play the role of several factors: habitat, habitat light conditions, lifestyles and evolutionary history.
Conclusion:

Findings

1. His eyes studied species of molluscs Arion rufus, Perforatella incarnata, Helicigona lapicida, Arianta arbustorum and Cepaea hortensis constructed according to the type of chamber with eyes fixed optics.

2. Dioptric apparatus consists of two lenses: the lens and the lens formed by the tentacular epidermis and cornea. In all studied species of molluscs are the dominant lenses optical components. Lenses A rufus, N. lapicida, A. arbustorum and C. hortensis, obviously, have a gradient of refractive index. The lens P. incarnata, apparently, is optically homogeneous.

3. Photosensitive device consists of two morphologically different types of photoreceptors mikrovillyarnyh. Do Ya lapicida photoreceptors of the second type are represented by three species, which probably means the presence of color vision in the mollusk. Svetoizoliruyuschy apparatus form the pigment cells that escape receptor cells from each other at the level of the cell bodies. Microvillus not optically isolated.

4. At A rufus, P. incarnata and I lapicida, with different lighting preferences, found an inverse relationship between the brightness of the preferred habitat, and optical sensitivity of the photoreceptors of the first type. At A arbustorum and C. hortensis, living under the same lighting conditions, found similar values ​​of the optical sensitivity of the photoreceptors of the first type. In all studied species of molluscs between sensitivity to light photoreceptors and brightness of the second type of habitat is no connection has been observed. Within the eyes of each type of shellfish seen the following relations: a high optical sensitivity of the photoreceptors of the first type / low optical sensitivity of the photoreceptors and the second type of low spatial resolution of the photoreceptor of the first type / high spatial resolution of the photoreceptor of the second type (with the exception of the second type of photoreceptor in the second type I lapicida). It may be more sensitive photoreceptors of the first type operate at low light levels, and the photoreceptors of the second type - with a bright light.

5. Features of the structure and optics of the eye A. rufus, P incarnata and I lapicida allow them as a function of the vision. Clams can be used for visual function realization of the basic forms of behavior: protective and food, as well as to detect konspetsificheskih individuals. Eye sensitivity A. rufus and P. incarnata will allow them these visual tasks in daytime and twilight, and J. lapicida - only in the daytime. Adaptive devices to the vision of the environment can be traced in two directions: one of them focused on achieving a relatively high visual acuity, others - to achieve adequate sensitivity to the available light. In the former case, adaptation can be expressed by the lengthening of the focal length of the optical system of the eye and / or increasing the density of photoreceptor cells. In the second case of adaptation appear to change the diameter of the aperture and the size (diameter and length) svetovosprinimayuschih parts of photoreceptor cells of the first type in accordance with the brightness of the preferred habitats of shellfish. A arbustorum eyes and C hortensis can function as light detectors and provide the possibility of finding the optimal habitats for life lighting conditions.

6. Major changes in the structure of the eye caused by the passage of mollusks in the phylogenetic development of the water in the terrestrial environment, can be traced in the direction of thickening of the cornea, the acquisition of a soft consistency and ellipsoid lenses, as well as reducing the distance between the lens and the retina to the minimum possible. Apparently, the ancestors of perednezhabernyh clams have inherited the dominant role of the lens in focusing the light and the gradient of refractive index of the lenses.

7. In the formation of the eyes of mollusks play the role of several factors: habitat, habitat light conditions, lifestyles and evolutionary history.

Conclusion

As already noted, during the evolution of certain groups of mollusks Class Gastropoda adapted to terrestrial life (Cook, 2001). From sea to land snails can get in different ways. Firstly, through the estuary - the broad mouth of the river are available for hot flashes - in fresh water such as lakes and rivers, and then - on dry land. Secondly, through the mangroves - trees and shrub plant communities developed in periodically flooded areas of the sea coasts and estuaries (Fechter, Falkner, 1990). Development of new habitat has caused the need for modifications to existing systems of organs, including the visual sensory system. The most striking difference between the majority of terrestrial molluscs (Stylommatophora) from mollusks living in the water (. Basommatophora), is the location of the eyes on the tops of the additional (upper) pair of tentacles, which has greatly expanded their visual field (Brehm, 1948; Cook, 2001). Some other special features listed below are also adapting peripheral part of the visual system to function on land.

Lenses in all investigated in this work mollusks noncellular, ie formed secretory material. They are surrounded by only the corneal cells and cells of the retina. Therefore we can assume that, as in other gastropods, the source of building material for lens A. rufus, P. incarnata, N. lapicida, A. and C. hortensis arbustorum are precisely the cells of the cornea and / or supporting cells of the retina. Formed lenses are elliptical shape and a relatively soft consistency, as well as many other lenses of the class Gastropoda, living in the terrestrial environment, such as snails, C. nemoralis and T. hispida (Shepeleva, 2005). These lenses are fully occupy the cavity of the eye, leaving the vitreous space of only a few microns. Lenses studied, the vast majority of aquatic (marine and freshwater), gastropods have opposite characteristics: a spherical or near-field shape, firm texture, gradient refractive index and the gap between the lens and retina mikrovillyarnym layer, which is equal to several tens or hundreds of microns. In these animals the complex epithelium / cornea lens shortens the focal length of only 1-3 microns. Therefore, the lens becomes not just a major, but in fact the only component of the refractive optics of the eye and, therefore, must be powerful. In this regard, there is a need in its spherical shape, firm texture and gradient of refractive index, which give the shortest focal length. And so the image falls on the retina, it must be separated from the lens, as focal length is measured from its center. In the studied terrestrial mollusks lens, although it remains the dominant optical element, the combination of the epithelium / cornea contributes significantly to the creation of images on the receptor layer of the retina, reducing the lens focal length by 40-70 mm. Focal length is relative to the rear of the main plane of the optical system, which does not coincide with the center of the lens. Therefore, land-shells do not need to maintain the same properties dioptric apparatus and the same principle of arrangement of its components. However, the gradient of refractive index lens material is clearly present in four of the five species studied. It should also be noted that the thickness of the cornea in the studied land-shells are several times greater than the thickness of the cornea of ​​aquatic species, which probably reflects the need for eye protection against loss of moisture in the air. With regard to the tentacular epidermis, no trends in the thickness of the terrestrial molluscs in comparison with water molluscs are not observed. Thus, of all the above it is clear that changes in eye diopter apparatus shellfish, have fallen into the phylogenetic development of the water to a terrestrial lifestyle, are adapting to the functioning of the new-land habitat. Major changes can be traced in the direction of thickening of the cornea, the acquisition of a soft consistency and ellipsoid lenses, as well as reducing the distance between the lens and the retina to the minimum possible. Apparently, from the anterior gill-ancestors have inherited not just clams dominant role in the lens focusing light and refractive index gradient material of the lenses.

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