Mechanisms of accommodation in the bird eye

Summary Keratoscopic study of corneal curvature before and after accommodation in two common bird species failed to provide evidence of a corneal accommodative mechanism. Accommodative changes in refractive state measured retinoscopically are presumably brought about by the effect of ciliary muscle contraction on lens curvature. However, retinoscopic and freeze-sectioning study of accommodation in diving ducks supports the long suspected existence of an iris accommodative mechanism capable of producing dramatic changes in lens curvature. This mechanism is believed to be a means of compensating for the refractive loss of the cornea in water.This research was supported in part by a grant from the Canadian National Sportsmen's Fund. The assistance of Dr. W.F. Long and Mr. R. Ortleib (Niska Wildlife Foundation) is gratefully acknowledged.     

Corneal accommodation in chick and pigeon

Summary We have investigated the role of changes in corneal radius of curvature in effecting accommodation in the bird's eye. It was found that in natural accommodation (measured by IR photoretinoscopy) changes of corneal radius of curvature (measured by IR photokeratometry) play an important role in both the chick and the pigeon. In the adult pigeon the cornea is indeed responsible for the largest part of natural accommodation (up to approx. 9 D). In this animal the corneal diameter (as seen from the optical axis of the eye) decreases in accommodation which can be taken to explain the change of corneal radius of curvature. In the chicken, corneal accommodation is combined with other mechanisms (total accommodative range 15–17 D, corneal accommodation about 8 D). The chicken's cornea is aspherical within the pupil area leading to large measurement variation in photokeratometry if the Purkinje images are not symmetrical to the pupillary axis.Abbreviation IR infrared     

The action of ciliary muscle contraction on accommodation of the lens explored with a 3D model

Biomechanics and Modeling in Mechanobiology - The eye’s accommodative mechanism changes optical power for near vision. In accommodation, ciliary muscle excursion relieves lens tension,...     

A comparative study of the adrenergic innervation of the iris and ciliary structures in 18 species in phylogenesis

A detailed topography of adrenergic innervation in invertebrates (lobster), low vertebrates (fish, amphibians, reptiles, birds), and nine species of mammals is presented. Flack and Hillarp's specific fluorescent histochemical method using freeze-dried material was used. Phylogenetically, adrenergic innervation appeared earlier under the ciliary epithelium and in the muscle than surrounding the vessels, and in all species many fibers were without any connection to the vessel walls. Adrenergic innervation was very rich in the dilator muscle extending toward the epithelium of the posterior chamber; a surprisingly rich network was found in the sphincter muscle and also in ciliary spaces of some species. Numerous fluorescent mast cells were visualized in the pecten of the bird eye and in the ciliary tissue of the sheep and cow.     

The evolution of the mechanical properties of amniote bone

162 specimens from 19 species of amniote were tested for various mechanical and physical properties to ascertain whether there were characteristic differences between different groups. All mechanical properties showed very great variation. In general the reptiles were not inferior to the mammals and birds. The histology of living forms was compared to that of fossil forms, to see whether 'weak' histology was more characteristic of primitive amniotes. The earliest reptiles probably had rather complaint bone, but it was probably tough. Modern types of bone appeared over two hundred million years ago. Very specialised bone, like that of the bullae of whales and antlers, may have evolved only in the mammals, but the fossil record is not complete enough to assert this confidently.     

Analysis of human crystalline lens accommodation

The behavior of the human crystalline lens during accommodation is analytically studied. The lens is modeled as a closed axisymmetrical membrane shell of varying thickness enclosing an incompressible liquid. To simulate zonular tension associated with lenticular accommodation, an axisymmetrical radial force or displacement is imposed around the shell equator. Two second-order, simultaneous, nonlinear governing differential equations are derived. Numerical results, obtained from the investigation of human lens profiles of three independently published MRI images and a drawing of a microphotograph, demonstrate that when zonular traction within the physiological force range of the ciliary muscle is exerted, both central lens thickness and central optical power increase. Qualitatively, these increases are independent of lens shape. However, the magnitude of these changes is dependent on the initial profile of the lens and is enhanced by the "natural" variation in capsular thickness. Only when a pulling force significantly exceeds the force capacity of the ciliary muscle does the lens flatten and its central thickness and optical power decrease.     

Some introductory notes on the organization of the forebrain in tetrapods.

As an introduction to the main theme of this conference an overview of the organization of the tetrapod forebrain is presented with emphasis on the telencephalic representation of sensory and motor functions. In all classes of tetrapods, olfactory, visual, octavolateral, somatosensory and gustatory information reaches the telencephalon. Major differences exist in the telencephalic targets of sensory information between amphibians and amniotes. In amphibians, three targets are found: the lateral pallium for olfactory input, the medial pallium for visual and multisensory input, and the lateral subpallium for visual, octavolateral and somatosensory information. The forebrains of reptiles and mammals are similar in that the dorsal surface of their cerebral hemisphere is formed by a pallium with three major segments: (a) an olfactory, lateral cortex; (b) a 'limbic' cortex that forms the dorsomedial wall of the hemisphere, and (c) an intermediate cortex that is composed entirely of isocortex in mammals, but in reptiles (and birds) consists of at least part of the dorsal cortex (in birds the Wulst) and a large intraventricular protrusion, i.e. the dorsal ventricular ridge. In birds, the entire lateral wall of the hemisphere is involved in this expansion. The intermediate pallial segment receives sensory projections from the thalamus and contains modality-specific sensory areas in reptiles, birds and mammals. The most important differences between the intermediate pallial segment of amniotes concern motor systems.     

Germline development in amniotes: A paradigm shift in primordial germ cell specification

In the field of germline development in amniote vertebrates, primordial germ cell (PGC) specification in birds and reptiles remains controversial. Avians are believed to adopt a predetermination or maternal specification mode of PGC formation, contrary to an inductive mode employed by mammals and, supposedly, reptiles. Here, we revisit and review some key aspects of PGC development that channelled the current subdivision, and challenge the position of birds and reptiles as well as the ‘binary’ evolutionary model of PGC development in vertebrates. We propose an alternative view on PGC specification where germ plasm plays a role in laying the foundation for the formation of PGC precursors (pPGC), but not necessarily of PGCs. Moreover, inductive mechanisms may be necessary for the transition from pPGCs to PGCs. Within this framework, the implementation of data from birds and reptiles could provide new insights on the evolution of PGC specification in amniotes.     

Volume change of the ocular lens during accommodation

During accommodation, mammalian lenses change shape from a rounder configuration (near focusing) to a flatter one (distance focusing). Thus the lens must have the capacity to change its volume, capsular surface area, or both. Because lens topology is similar to a torus, we developed an approach that allows volume determination from the lens cross-sectional area (CSA). The CSA was obtained from photographs taken perpendicularly to the lenticular anterior-posterior (A-P) axis and computed with software. We calculated the volume of isolated bovine lenses in conditions simulating accommodation by forcing shape changes with a custom-built stretching device in which the ciliary body-zonulae-lens complex (CB-Z-L) was placed. Two measurements were taken (CSA and center of mass) to calculate volume. Mechanically stretching the CB-Z-L increased the equatorial length and decreased the A-P length, CSA, and lens volume. The control parameters were restored when the lenses were stretched and relaxed in an aqueous physiological solution, but not when submerged in oil, a condition with which fluid leaves the lens and does not reenter. This suggests that changes in lens CSA previously observed in humans could have resulted from fluid movement out of the lens. Thus accommodation may involve changes not only in capsular surface but also in volume. Furthermore, we calculated theoretical volume changes during accommodation in models of human lenses using published structural parameters. In conclusion, we suggest that impediments to fluid flow between the aquaporin-rich lens fibers and the lens surface could contribute to the aging-related loss of accommodative power. lens volume calculation; intralenticular fluid movement; presbyopia; mammalian lens     

Cerebral cortical and brainstem areas related to the central control of lens accommodation in cat and monkey.

1. Studies on the central nervous system related to lens accommodation in cat and monkey were reviewed. 2. During the last decade, a considerable amount of neurophysiological data on the peripheral innervation of the ciliary muscle, properties of parasympathetic oculomotor neurons and mesencephalic reticular neurons have accumulated. 3. Neurophysiological and anatomical evidence supporting the contribution of the cerebellum to lens accommodation has been obtained. 4. Recently, cerebral cortical neurons in the parieto-occipital cortex with activities related to lens accommodation were found in cat and monkey.     

Neurochemical organization of reptilian thalamus. Comparative analysis of amniote optical centers

Study of neurochemical characteristics of the turtle thalamus was performed using antibodies against several biologically active compounds; monoamines (5-HT, TH), neuropeptides (SP, m-Enk, NPY), against ChAT and by histochemical detection of NADPH-d. Based on our own results and literature data obtained on other representatives of reptiles, birds, and mammals, a comparative analysis was carried out of neurochemical organization of two thalamic optical centers—relay nuclei of thalamofugal (GLd) and tectofugal (Rot/LP-Pulv) systems of amniotes. Features of similarities and differences of these centers in representatives of non-mammalian and mammalian amniotes are revealed. GLd has a great similarity of the studied neurochemical characteristics in all amniotes. It receives innervation from all studied transmitter-modulatory systems with predominance of serotonin-, choline- and NPY-ergic projections. Neurochemical organization of Rot comparable with the tectorecipient part of mammalian LP-Pulv has a great resemblance in reptiles and birds, with considerable interspecies differences inside each class. In the majority of the studied systems, Rot is characterized by scant innervation. On the contrary, LP-Pulv receives sufficiently massive innervation from these systems. The most characteristic of this complex are rich SP- and cholinergic projections that are scant in Rot of reptiles and birds. The similar feature of Rot and LP-Pulv is the presence of massive serotonin- and NO-ergic (NADPH-positive) projections. The revealed similarities and differences of the neurochemical characteristics of thalamic optical centers among amniotes seem to reflect various transformations of thalamo- and tectofugal visual systems in the course of phylogenetic and adaptive evolution.     

The relationship between body mass and relative investment in testes mass in amniotes and other vertebrates

In terrestrial placental mammals, there is a well‐known negative allometric relationship between body mass and relative investment in testes mass. Such a negative relationship means that males of relatively monogamous small species invest proportionately more in their reproductive tissues than males of more polyandrous larger species. The selective pressure responsible for this relationship remains unclear and is it not known if this is a general allometric relationship that is similar across all vertebrate lineages. To investigate this, we conducted the first comparison of relationships between body mass and testes mass (using percentage testes mass as the dependent variable) across a variety of vertebrate groups. In all amniote lineages examined, the allometric relationship between body mass and testes mass was relatively strong and negative. We show, for the first time, that reptiles, birds and terrestrial placental mammals followed the same allometric relationship and, contrary to previous expectations, this relationship is sigmoidal rather than linear. Within this data set, there was no significant difference between this general amniote relationship and any of the 13 orders of reptiles, birds and terrestrial placental mammals examined. As a result, we propose that a sigmoidal relationship should be considered the default assumption for the form of the body mass – testes mass relationship within the amniote lineage. However, we also identify significant differences within some additional mammal groups (marsupials, bats and cetaceans). In each of these cases, only some sub‐groupings differed significantly from the general amniote relationship. In contrast to the amniotes, the relationship is relatively weak and positive in teleost fish and frogs suggesting that a negative allometric relationship is not universal in vertebrates. We explore whether variation in the body mass – testes mass relationships can be linked to sperm competition or a variety of ecological characteristics, either for amniotes in particular or vertebrates in general.     

Immunofluorescence study of the formation of evolutionary stable lens proteins in chick embryos]

In the lens of fishes (carp, spiny dogfish) beta-crystallins were identified which were characteristic also of reptiles, amphibians, birds and mammals (evolutionary stable beta-crystallins). The dynamics of the formation of such beta-crystallins in 5--14 days old chick embryos was studied by the indirect immunofluorescence method with antisera to fish lens. These proteins are reliably indentified first at the lens sections from 7--8days old chick embryos. At all stages under study these beta-crystallins are localized mainly in the epithelial cells and practically not found in the lens fibers. They were, however, found in the fibrous (central) part of developing lens as well by the method of immunoelectrophoresis.     

An exploration of differences in the scaling of life history traits with body mass within reptiles and between amniotes

Allometric relationships linking species characteristics to body size or mass (scaling) are important in biology. However, studies on the scaling of life history traits in the reptiles (the nonavian Reptilia) are rather scarce, especially for the clades Crocodilia, Testudines, and Rhynchocephalia (single extant species, the tuatara). Previous studies on the scaling of reptilian life history traits indicated that they differ from those seen in the other amniotes (mammals and birds), but so far most comparative studies used small species samples and also not phylogenetically informed analyses. Here, we analyzed the scaling of nine life history traits with adult body mass for crocodiles (n = 22), squamates (n = 294), turtles (n = 52), and reptiles (n = 369). We used for the first time a phylogenetically informed approach for crocodiles, turtles, and the whole group of reptiles. We explored differences in scaling relationships between the reptilian clades Crocodilia, Squamata, and Testudines as well as differences between reptiles, mammals, and birds. Finally, we applied our scaling relationships, in order to gain new insights into the degree of the exceptionality of the tuatara's life history within reptiles. We observed for none of the life history traits studied any difference in their scaling with body mass between squamates, crocodiles, and turtles, except for clutch size and egg weight showing small differences between these groups. Compared to birds and mammals, scaling relationships of reptiles were similar for time‐related traits, but they differed for reproductive traits. The tuatara's life history is more similar to that of a similar‐sized turtle or crocodile than to a squamate.     

The amino acid sequences of two alpha chains of hemoglobins from Komodo dragon Varanus komodoensis and phylogenetic relationships of amniotes

To elucidate phylogenetic relationships among amniotes and the evolution of alpha globins, hemoglobins were analyzed from the Komodo dragon (Komodo monitor lizard) Varanus komodoensis, the world's largest extant lizard, inhabiting Komodo Islands, Indonesia. Four unique globin chains (alpha A, alpha D, beta B, and beta C) were isolated in an equal molar ratio by high performance liquid chromatography from the hemolysate. The amino acid sequences of two alpha chains were determined. The alpha D chain has a glutamine at E7 as does an alpha chain of a snake, Liophis miliaris, but the alpha A chain has a histidine at E7 like the majority of hemoglobins. Phylogenetic analyses of 19 globins including two alpha chains of Komodo dragon and ones from representative amniotes showed the following results: (1) The a chains of squamates (snakes and lizards), which have a glutamine at E7, are clustered with the embryonic alpha globin family, which typically includes the alpha D chain from birds; (2) birds form a sister group with other reptiles but not with mammals; (3) the genes for embryonic and adult types of alpha globins were possibly produced by duplication of the ancestral alpha gene before ancestral amniotes diverged, indicating that each of the present amniotes might carry descendants of the two types of alpha globin genes; (4) squamates first split off from the ancestor of other reptiles and birds.      

Focusing and accommodation in the brown kiwi (Apteryx australis)

Summary Brown kiwis are an endangered species of nocturnal, flightless birds which are native to New Zealand. The resting focus of two specimens has previously been studied by retinoscopy in a zoo while the birds were restrained by their keeper (Sivak and Howland 1987). Those birds appeared to be hyperopic (farsighted) by 2–7 D. In this study, examination with infrared photorefraction of the focusing of two unrestrained, feeding birds showed that they could focus objects at infinity and objects in their immediate environment and that they had modest powers of accommodation. Measurements on two 6 month old kiwi chicks showed their corneal radius of curvature to be between 2.90 and 3.00 mm (117 D and 101 D in power).     

Focusing and accommodation in tuatara (Sphenodon punctatus)

Summary Photorefraction and photokeratometry were performed on two juvenile tuatara (Sphenodon punctatus, 7 years of age, total length approx. 27 cm). Sphenodon is the only surviving genus of the Rhynchocephalia, an order of reptiles. Both existing species are endangered and are found only in New Zealand. Even though Walls (1942) has stated that the ciliary muscle is relatively weak in tuataras, we observed 8 D of accommodation. The eyes were found to focus independently and we could induce an anisometropia of 6 D. The average corneal power of the tuatara was found to be 101 D.     

The origin of homoiothermy--unsolved problem

The analysis of allometric dependence of energy expenditure on body mass among reptiles, birds and mammals has shown that standard metabolic rate of reptiles when they are warmed up to the temperature of homoiothermic animals is an order of magnitude lower than that of birds and mammals. Basal metabolism is originated as special feature historically related to the metabolism during active behavior, rather than thermal regulation. Facultative endothermy was not advantageous for large animals because of long time needed to warm up the body. The ancestors of birds and animals escaped negative consequences of van't-Hoff equation by choosing constant body temperature. Heat conductivity of reptile's covers is so great, that it cannot keep endogenous warm of resting animal at any temperature of the body. Reptile "dressed" in covers of bird or mammal would be able to keep warm under conditions of maximal aerobic muscular activity and body temperature similar to that of homoiothermic animals. The base of chemical thermoregulation in birds and mammals is a thermoregulatory muscle tonus which remains unknown. One can suppose that during evolution of birds and mammals the saltation-liked origin of endothermy "fixed" the level of metabolism typical for running reptile and transformed in into the basal metabolism. This event took place at the cell and tissue level. The absence of palaeontological evidences and intermediate forms among recent species does not allow easy understanding of homoiothermy origin.     

Adaptation to the land: The skin of reptiles in comparison to that of amphibians and endotherm amniotes

The adaptation to land from amphibians to amniotes was accompanied by drastic changes of the integument, some of which might be reconstructed by studying the formation of the stratum corneum during embryogenesis. As the first amniotes were reptiles, the present review focuses on past and recent information on the evolution of reptilian epidermis and the stratum corneum. We aim to generalize the discussion on the evolution of the skin in amniotes. Corneous cell envelopes were absent in fish, and first appeared in adult amphibian epidermis. Stem reptiles evolved a multilayered stratum corneum based on a programmed cell death, intensified the production of matrix proteins (e.g., HRPs), corneous cell envelope proteins (e.g., loricrine-like, sciellin-like, and transglutaminase), and complex lipids to limit water loss. Other proteins were later produced in association to the soft or hairy epidermis in therapsids (e.g., involucrin, profilaggrin-filaggrin, trichohyalin, trichocytic keratins), or to the hard keratin of hairs, quills, horns, claws (e.g., tyrosine-rich, glycine-rich, sulphur-rich matrix proteins). In sauropsids special proteins associated to hard keratinization in scales (e.g., scale beta-keratins, cytokeratin associated proteins) or feathers (feather beta-keratins and HRPs) were originated. The temporal deposition of beta-keratin in lepidosaurian reptiles originated a vertical stratified epidermis and an intraepidermal shedding layer. The evolutions of the horny layer in Therapsids (mammals) and Saurospids (reptiles and birds) are discussed. The study of the molecules involved in the dermo-epidermal interactions in reptilian skin and the molecular biology of epidermal proteins are among the most urgent future areas of research in the biology of reptilian skin.     

Cytochemical, biochemical and molecular aspects of the process of keratinization in the epidermis of reptilian scales

The characteristics of scaled skin of reptiles is one of their main features that distinguish them from the other amniotes, birds and mammals. The different scale patterns observed in extant reptiles result from a long evolutive history that allowed each species to adapt to its specific environment. The present review deals with comparative aspects of epidermal keratinization in reptiles, chelonians (turtles and tortoises), lepidosaurian (lizards, snakes, sphenodontids), archosaurians (crocodilians). Initially the morphology and cytology of reptilian scales is outlined to show the diversity in the epidermis among different groups. The structural proteins (alpha-keratins and associated proteins), and enzymes utilized to form the corneous layer of the epidermis are presented. Aside cytokeratins (alpha-keratins), used for making the cytoskeleton, reptilian alpha-keratinocytes produce interkeratin (matrix) and corneous cell envelope proteins. Keratin bundles and degraded cell organelles constitute most of the corneous material of alpha-keratinocytes. Matrix, histidine-rich and sulfur-rich proteins are produced in the soft epidermis and accumulated in the cornified cell envelope. Main emphasis is given to the composition and to the evolution of the hard keratins (beta-keratins). Beta-keratins constitute the hard corneous material of scales. These small proteins are synthesized in beta-keratinocytes and are accumulated into small packets that rapidly merge into a compact corneous material and form densely cornified layers. Beta-keratins are smaller proteins (8-20 kDa) in comparison to alpha-keratins (40-70 kDa), and this size may determine their dense packing in corneocytes. Both glycine-sulfur-rich and glycine-proline-rich proteins have been so far sequenced in the corneous material of scales in few reptilian species. The latter keratins possess C- and N-amino terminal amino acid regions with sequence homology with those of mammalian hard keratins. Also, reptilian beta-keratins possess a central core with homology with avian scale/feather keratins. Multiple genes code for these proteins and their discovery and sequentiation is presently an active field of research. These initial findings however suggest that ancient reptiles already possessed some common genes that have later diversified to produce the specific keratin-associated proteins in their descendants: extant reptiles, birds and mammals. The evolution of these small proteins in lepidosaurians, chelonians and archosaurians represent the next step to understand the evolution of cornification in reptiles and derived amniotes (birds and mammals).