A model for the temporal organization of X- and Y-type receptive fields in the primate retina

A model is proposed for the temporal characteristics of X-and Y-type responses of ganglion cells in the primate retina. The main suggestions of the model are: (I) The X-type temporal response is determined primarily by the delay between center and surround contributions. (II) The Y-type response is generated in the inner plexiform layer by a derivativelike operation on the bipolar cell's input, followed by a rectification in the convergence of these inputs onto the Y-ganglion-cell. (III) The derivative-like operation is obtained by recurrent inhibition in the dyad synaptic structure.The X-and Y-type responses predicted by the model, for a variety of stimuli, were examined and compared with available electrophysiological recordings. Finally, certain predictions derived from the model are discussed.     

Receptive field mechanisms of ganglion cells in the cat retina

On the basis of anatomical and physiological results of the vertebrate retina, a method is proposed for analysing the respective fields of ganglion cells in the cat retina. In the model, we assume the following: (a) Ganglion cells receive their input from bipolar and/or amacrine cells. (b) The nonlinearity of ganglion cell responses is due to the activities of transient type amacrine cells. The method has been proved to be effective. According to the results of this investigation, the receptive field properties of X type and Y type ganglion cells are heterogeneous. Thus, it may be considered that their receptive fields consist of center and surround mechanisms. The receptive field properties of X-cells are almost linear and the X-cells seem to receive most of their input from bipolar cells. On the other hand, the ones of Y-cells are highly nonlinear. Consequently, it is conceivable that the Y-cells receive their input mainly from transient type amacrine cells.     

A unified neural model of spatiotemporal processing in X and Y retinal ganglion cells

This work presents unified analyses of spatial and temporal visual information processing in a feed-forward network of neurons that obey membrane, or shunting equations. The feed-forward shunting network possesses properties that make it well suited for processing of static, spatial information. However, it is shown here that those same properties of the shunting network that lead to good spatial processing imply poor temporal processing characteristics. This article presents an extension of the feed-forward shunting network model that solves this problem by means of preprocessing layers. The anatomical interpretation of the resulting model is structurally analogous to recently discovered data on a retinal circuit connecting cones to retinal ganglion cells through pairs of pushpull bipolar cells. Mathematical analysis of the lumped model leads to the hypothesis that X and Y retinal ganglion cells may consist of a single mechanism acting in different parameter ranges. This hypothesis is confirmed in the companion article, wherein the model in conjunction with a nonlinear temporal adaptation mechanism — is used to reproduce experimental data of both X and Y cells by simple changes in morphological and physiological parameters.     

A spatio-temporal model of ganglion cell receptive field in the cat retina

A spatio-temporal model of ganglion cell receptive fields is proposed on the basis of receptive field characteristics of cat retinal ganglion cells reported in our previous paper. The model consists of the linear and nonlinear mechanisms in the ganglion cell receptive field. The linear mechanism is assumed to be composed of antagonistic center and surround mechanisms. Then, by integrating these mechanisms we construct a spatio-temporal impulse response function of ganglion cell receptive field. Here we assume that spatio-temporal impulse response function may be factored into spatial and temporal terms. By Fouriertransforming the spatio-temporal impulse response function, we can obtain the spatio-temporal transfer function. Contrast sensitivity characteristics of X-and Y-cells in the cat retina may be explained by the transfer function.     

Optimization of polylactic-co-glycolic acid nanoparticles containing itraconazole using 2<Superscript>3</Superscript> factorial design

This study investigated the utility of a 2³ factorial design and optimization process for polylactic-co-glycolic acid (PLGA) nanoparticles containing itraconazole with 5 replicates at the center of the design. Nanoparticles were prepared by solvent displacement technique with PLGAX ₁ (10, 100 mg/mL), benzyl benzoateX ₂ (5, 20 μg/mL), and itraconazoleX ₃ (200, 1800 μg/mL). Particle size (Y ₁), the amount of itraconazole entrapped in the nanoparticles (Y ₂), and encapsulation efficiency (Y ₃) were used as responses. A validated statistical model having significant coefficient figures (P<.001) for the particle size (Y ₁), the amount of itraconazole entrapped in the nanoparticles (Y ₂), and encapsulation efficiency (Y ₃) as function of the PLGA (X ₁), benzyl benzoate (X ₂), and itraconazole (X ₃) were developed: Y₁=373.75+66.54X₁+52.09X₂+105.06X₃−4.73X₁X₂+46.30X₁X₃; Y₂=472.93+73.45X₁+ 169.06X₂+333.03X₃+62.40X₁X₃+141.49X₂X₃; Y₃= 57.36+6.53X₁+15.52X₂−12.59X₃+1.01X₁X₃+ 1.73X₂X₃.X ₁,X ₂, andX ₃ had a significant effect (P<.001) onY ₁,Y ₂, andY ₃. The particle size, the amount of itraconazole entrapped in the nanoparticles, and the encapsulation efficiency of the 4 formulas were in agreement with the predictions obtained from the models (P<.05). An overlay plot for the 3 responses shows the boundary in whichY ₁ shows the boundary in which a number of combinations of concentration of PLGA, benzyl benzoate, and itraconazole will result in a satisfactory process. Using the desirability approach with the same constraints, the solution composition having the highest overall desirability (D=0.769) was 10 mg/mL of PLGA, 16.94 μg/mL of benzyl benzoate, and 1001.01 μg/mL of itraconazole. This approach allowed the selection of the optimum formulation ingredients for PLGA nanoparticles containing itraconazole of 500 μg/mL.     

P2Y receptors and pain transmission

It is widely accepted that the most important ATP receptors involved in pain transmission belong to the P2X₃ and P2X_2/3 subtypes, selectively expressed in small diameter dorsal root ganglion (DRG) neurons. However, several types of the metabotropic ATP (P2Y) receptors have also been found in primary afferent neurons; P2Y₁ and P2Y₂ receptors are typically expressed in small, nociceptive cells. Here we review the results available on the involvement of P2Y receptors in the modulation of pain transmission.     

Characterisation of columnar neurons and visual signal processing in the medulla of the locust optic lobe by system identification techniques

We describe visual responses of seventeen physiological classes of columnar neuron from the retina, lamina and medulla of the locust (Locusta migratoria) optic lobe. Many of these neurons were anatomically identified by neurobiotin injection. Characterisation of neuronal responses was made by moving and flash stimuli, and by two system identification techniques: 1. The first-order spatiotemporal kernel was estimated from response to a spatiotemporal white-noise stimulus; 2. A set of kernels to second order was derived by the maximal-length shift register (M-sequence) technique, describing the system response to a two-channel centre-surround stimulus. Most cells have small receptive fields, usually with a centre diameter of about 1.5°, which is similar to that of a single receptor in the compound eye. Linear response components show varying spatial and temporal tuning, although lateral inhibition is generally fairly weak. Second-order nonlinearities often have a simple form consistent with a static nonlinear transformation of the input from the large monopolar cells of the lamina followed by further linear filtering.Abbreviations LMC large monopolar cell - LVF long visual fibre - RF receptive field - SMC small monopolar cell - SVF short visual fibre     

Initial processing of visual information within the retina and the LGN

The initial stage of information processing by the visual system reduces the information contained in the continuous image on the retina into a discrete set of responses which are carried from the lateral geniculate nucleus (LGN) to the visual cortex.-1. The optimal sampling of the light intensity distribution in the visual environment is achieved only if each channel in the visual pathways carries undistorted information corresponding to an image element. The visual system approaches as closely as possible the scheme of optimal spatial sampling, retaining the full information on the low spatial frequency content of the object light intensity. The ideal receptive field of a sustained LGN cell is then of the form J ₁ (Kr)/Kr.-2. The experimentally determined receptive fields of sustained LGN cells (and to some extent retinal ganglion cells as well) in cat closely resemble the functional form J ₁ (Kr)/Kr. The centre-surround organization of the receptive fields is therefore understood as a scheme which leads to a maximal information flow through the visual pathways.-3. The optimal sampling scheme cannot be realized by the retina alone, because of restrictions on the size of neural networks. It is therefore constructed in two stages, ending at the LGN level. A recombination of ganglion cell signals into optimal receptive fields is a major role of the LGN.     

The Mean of the Inverse of a Punctured Normal Distribution and Its Application

The fundamental properties of a punctured normal distribution are studied. The results are applied to three issues concerning X/Y where X and Y are independent normal random variables with means μ_X and μ_Y respectively. First, estimation of μ_X/μ_Y as a surrogate for E(X/Y) is justified, then the reason for preference of a weighted average, over an arithmetic average, as an estimator of μ_X/μ_Y is given. Finally, an approximate confidence interval for μ_X/μ_Y is provided. A grain yield data set is used to illustrate the results. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Sex determination of the onion fly,Hylemya antiqua (Meigen)

X and Y chromosomes of Hylemya antiqua occur in two forms each. X_L and X_S, and Y₁ and Y₂. The larger X_L has an intercalary proximal segment which is absent in the more common smaller X_S. The acrocentric Y chromosome (Y₁), does not differ morphologically from X_S. A smaller metacentric Y₂ is apparently not homologous with Y₁. Two types of males, XY₁ and XXY₂, coexist in at least one Dutch population. XY₂ has been observed in one individual only. In larval ganglion cells an association has been observed between chromosome Y₂ and a probably non-homologous, intercalary segment of autosome 4. A numerical somatic variation of Y₂ can lead to gynandromorphs and sex ratios significantly different from 1∶1. XX cells can differentiate into functional spermatozoa in XX/XXY₂ mosaic testes. This indicates the presence of a diffusable male determining substance, which can reverse the “genotypic” sex of a cell. The occurrence of some spermatozoa-containing “cysts” in ovaries of two gynanders suggests a more or less autonomous (independent of the gonadal environment) differentiation of XXY₂ germ cells. XXXY₂ males and XXX females do not show a serious reduction in fertility. Even XXXXY₂ males do not exhibit any sign of intersexuality and spermatogenesis seems unaffected. All 62 scored M II cells of X-tetrasomic males contained 2 Xs.     

Two light‐activated neuroendocrine circuits arising in the eye trigger physiological and morphological pigmentation

Two biological processes regulate light‐induced skin colour change. A fast ‘physiological pigmentation change’ (i.e. circadian variations or camouflage) involves alterations in the distribution of pigment containing granules in the cytoplasm of chromatophores, while a slower ‘morphological pigmentation change’ (i.e. seasonal variations) entails changes in the number of pigment cells or pigment type. Although linked processes, the neuroendocrine coordination triggering each response remains largely obscure. By evaluating both events in Xenopus laevis embryos, we show that morphological pigmentation initiates by inhibiting the activity of the classical retinal ganglion cells. Morphological pigmentation is always accompanied by physiological pigmentation, and a melatonin receptor antagonist prevents both responses. Physiological pigmentation also initiates in the eye, but with repression of melanopsin‐expressing retinal ganglion cell activity that leads to secretion of alpha‐melanocyte‐stimulating hormone (α‐MSH). Our findings suggest a model in which eye photoperception links physiological and morphological pigmentation by altering α‐MSH and melatonin production, respectively.     

Localisation of P2Y<Subscript>1</Subscript> and P2Y<Subscript>4</Subscript> receptors in dorsal root,nodose and trigeminal ganglia of the rat

The presence and distribution of P2Y (nucleotide) receptor subtypes in rat sensory neurons has been investigated. RT-PCR showed that P2Y₁, P2Y₂, P2Y₄ and P2Y₆ receptor mRNA is expressed in sensory ganglia [dorsal root ganglion (DRG), nodose ganglion (NG) and trigeminal ganglion (TG)]. The regional and cellular distribution of P2Y₁ and P2Y₄ receptor proteins in these ganglia was investigated using immunohistochemistry. P2Y₁ polyclonal antibodies stained over 80% of the sensory neurons, particularly the small-diameter (neurofilament-negative) neurons. The P2Y₄ receptor antibody stained more medium- and large- (neurofilament-positive) diameter neurons than small-diameter neurons. P2Y₁ and P2Y₄ receptor immunoreactivity (P2Y₁-IR and P2Y₄-IR) was often coexpressed with P2X₃ receptor immunoreactivity (P2X₃-IR) in subpopulations of neurons. Double immunohistochemistry showed that 73–84% of P2X₃ receptor-positive neurons also stained for the P2Y₁ receptor in DRG, TG and NG while only 25–35% also stained for the P2Y₄ receptor. Subpopulations of P2Y₁-IR neurons were coexpressed with NF200, CGRP and IB₄; most P2Y₄-IR neurons were coexpressed with NF200, while only a few neurons were coexpressed with CGRP (10–20%) or with IB₄ (1–2%). The results suggest that P2Y as well as P2X receptor subtypes contribute to purinergic signalling in sensory ganglia.     

Statistical features of impulse trains in cat's lateral geniculate neurons

The spontaneous discharges which recorded extracellularly from cells in the lateral geniculate nucleus (LGN) of a cat were classified into the following 3 main groups depending upon the shapes of their interval histograms and autocorrelation functions: the gamma type whose interval histogram is fitted by a gamma distribution function and whose autocorrelation function has some periodic property which damps down within about several 10 ms, the burst type whose interval histogram has a peak in the first bin (less than 8 ms) and whose autocorrelation function has a large positive peak within several msec, and the multimodal type whose interval histogram has a complex shape with three or more peaks and whose autocorrelation function has a periodic property. Each type of spontaneous discharge seems to be inherent at scotopic and mesopic backgrounds, and the cells whose spontaneous discharges are the gamma type, the burst type, and the multimodal type are called here a gamma cell, burst cell, and the multimodal cell, respectively. Gamma cells are subdivided into X- and Y-cells (gamma-X and gamma-Y cells), but burst cells are all Y-cells and multimodal cells observed up to now are all X-cells. It is clear that these various types of cells are distributed significantly differently in each lamina. All the cells that we found up to now in lamina A were either burst cells or multimodal cells, but every type of cell was found in lamina A1. The majority of cells in lamina C were the gamma type. In most cases, the peak values of the PST histograms of gamma-Y cells (especially, on-center cells) are larger than those of burst cells. These results suggest that Y-cells projecting to area 17 from laminae A and A1 are the burst type, and Y-cells projecting to area 18 from laminae C and A1 are the gamma-Y type.     

The P2X7 receptor in retinal ganglion cells: A neuronal model of pressure-induced damage and protection by a shifting purinergic balance

Retinal ganglion cells process the visual signal and transmit it along their axons in the optic nerve to the brain. Molecular, immunohistochemical, and functional analyses indicate that the majority of retinal ganglion cells express the ionotropic P2X₇ receptor. Stimulation of the receptor can lead to a rise in intracellular calcium and cell death, although death does not involve the opening of a large diameter pore. Adenosine acting at A₃ receptors can attenuate the rise in calcium and death accompanying P2X₇ receptor activation, suggesting that dephosphorylation of ATP into adenosine is neuroprotective and that the balance of extracellular purines can influence neuronal survival. Increased intraocular pressure can lead to release of excessive extracellular ATP in the retina and damage ganglion cells by acting on P2X₇ receptors, implicating a role for the receptor in the loss of ganglion cell activity in glaucoma. In summary, the activation of P2X₇ receptors has both physiologic and pathophysiologic implications for ganglion cell function. These characteristics may also provide an insight into the contributions the P2X₇ receptor makes to neurons elsewhere.     

Properties of the sex chromosomes of Lucilia cuprina deduced from radiation studies

From a study of radiation-induced X-chromosome deletions the locus of black body (b) has been localized to the proximal portion of C-band defined euchromatin. Radiation produced mostly X-chromosome deletions rather than point mutations, total X or Y chromosome loss through breakage, or increased frequency of non-disjunction. Aberrant sex ratios obtained indicate that the X chromosome carries vital loci that were deleted with b ⁺ in many cases. The X/O karyotype produces fertile adult females with a characteristic phenotype which is also produced by X deletions. Sex chromosome non-disjunction to give X/O females and X/X/Y males is normally rare but is enhanced by the presence of chromosome rearrangements even when the X and Y are not involved.     

Purinergic receptors are part of a signalling system for proliferation and differentiation in distinct cell lineages in human anagen hair follicles

We investigated the expression of P2X₅, P2X₇, P2Y₁ and P2Y₂ receptor subtypes in adult human anagen hair follicles and in relation to markers of proliferation [proliferating cell nuclear antigen (PCNA) and Ki-67], keratinocyte differentiation (involucrin) and apoptosis (anticaspase-3). Using immunohistochemistry, we showed that P2X₅, P2Y₁ and P2Y₂ receptors were expressed in spatially distinct zones of the anagen hair follicle: P2Y₁ receptors in the outer root sheath and bulb, P2X₅ receptors in the inner and outer root sheaths and medulla and P2Y₂ receptors in living cells at the edge of the cortex/medulla. P2X₇ receptors were not expressed. Colocalisation experiments suggested different functional roles for these receptors: P2Y₁ receptors were associated with bulb and outer root sheath keratinocyte proliferation, P2X₅ receptors were associated with differentiation of cells of the medulla and inner root sheaths and P2Y₂ receptors were associated with early differentiated cells in the cortex/medulla that contribute to the formation of the hair shaft. The therapeutic potential of purinergic agonists and antagonists for controlling hair growth is discussed.     

Theoretical study on the aromaticity from d-AOs in cationic X 3 + (X = Sc,Y, La) clusters

The stable structures and aromatic characters for three cationic X₃⁺ (X = Sc, Y, and La) and three relevant neutral X₃Cl (X = Sc, Y, La) clusters are investigated at the DFT and post HF level of theory. The calculated results show that the X₃⁺ cations each has two stable structures: the regular trigon (D_3h) and the line (D_￥h {{\hbox{D}}_{\infty {\rm{h}}}} ) with the regular trigon (D_3h) being the ground state, while for three neutral X₃Cl clusters, Sc₃Cl has three stable isomers: the trigon-pyramidal (C_3v), bidentate (C_2v-1), and C_2v-2 structures, Y₃Cl and La₃Cl each has only two stable isomers: the trigon-pyramidal (C_3v) and bidentate (C_2v-1) structures. The ground states for three X₃Cl species are all the bidentate (C_2v-1) isomers. The calculations of the resonance energy (RE) and NICS show that trigonal X₃⁺ isomers exhibit higher degree of aromaticity. The detailed molecular orbital analyzes reveal that the isolated trigonal Sc₃⁺ and Y₃⁺ cations each has one delocalized π-type MO and shows single π-aromaticity, while the isolated trigonal La₃⁺ cation has one delocalized σ-type MO and shows single σ-aromaticity. The single π- or σ-aromaticity for X₃⁺ are attributed to the contributions mainly from the d AOs of the corresponding transition metal X atoms. However, when a singly negatively charged counterion Cl^- is added to Sc₃⁺, Y₃⁺, and La₃⁺ cations respectively, the aromatic type for the two Sc₃⁺, Y₃⁺ units in the corresponding neutral Sc₃Cl, Y₃Cl complexes are changed from π-aromaticity into σ-aromaticity, whereas the σ-aromaticity of the La₃⁺ units in the La₃Cl complex keeps unchanged in this process. Thus three Sc₃⁺, Y₃⁺, La₃⁺ units in the corresponding X₃Cl complexes all have only one σ-type MO and exhibit single σ-aromaticity.     

An example of a regular inbreeding system with cubic ancestral growth that preserves some genetic variability

A specific regular inbreeding system of quadruple half-second cousin mating is considered. A regular inbreeding system can be thought of as a graph which satisfies certain natural homogeneity properties. Random walks X _n and Y _n are introduced on the nodes of the graph; the event {X _n=Y_n} is a renewal event by the homogeneity properties. In Arzberger (1985) it is shown that 1) graphs associated with left cancellative semigroups are regular, and 2) for regular systems, the population becomes genetically uniform if and only if the event {X _n=Y_n} is recurrent. In Arzberger (1986) the system of quadruple half-second cousin mating is associated with a cancellative semigroup, thus the system is regular. In this paper we show that 1) A_n is asymptotically of the form cn ³, where A _n is the number of ancestors n generations into the past, 2) {X _n=Y_n} is not recurrent (this is shown by associating (X _n, Y _n) with a random walk in Z ³, 3) P[X _3n =Y _3n ] is asymptotically of the form cn ^–3/2. Thus, in this example, genetic heterogeneity is maintained, with a cubic rate of growth for A_n, not by an exponential growth rate, as in all previous examples of regular inbreeding systems in which genetic heterogeneity is maintained.     

Cytogenetic,hybridization and molecular evidence of four cytological forms of Anopheles nigerrimus (Hyrcanus Group) in Thailand and Cambodia

Thirteen isoline colonies of Anopheles nigerrimus were established from individual wild‐caught females collected from cow‐baited traps at locations in Thailand and Cambodia. Three types of X (X₁, X₂, X₃) and 4 types of Y (Y₁, Y₂, Y₃, Y₄) chromosomes were recovered, according to differing amounts of extra heterochromatin. Four karyotypic forms were designed depending upon apparently distinct figures of X and Y chromosomes, i.e., Form A (X₁, X₂, X₃, Y₁), B (X₂, X₃, Y₂), C (X₁, Y₃), and D (X₃, Y₄). Forms C and D were new metaphase karyotypes discovered in this study. Form A appeared to be common in both Thailand and Cambodia. Forms B and D were found to be rather specific to southern and northeastern Thailand, respectively, whereas Form C was confined to Cambodia. Hybridization experiments among the eight isoline colonies, which were representative of four karyotypic forms of An. nigerrimus, demonstrated genetic compatibility in giving viable progenies and synaptic salivary gland polytene chromosomes through F₂‐generations. These results elucidated the conspecific relationship, comprising four cytological forms within this taxon. Supportive evidence was confirmed further by very low intraspecific sequence variations (average genetic distance = 0.002–0.007) of the nucleotide sequences in ribosomal DNA [second internal transcribed spacer (ITS2)] and mitochondrial DNA [cytochrome c oxidase subunit I (COI) and subunit II (COII)].