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Texture segregation, surface representation and figure-ground separation.

A widespread view is that most texture segregation can be accounted for by differences in the spatial frequency content of texture regions. Evidence from both psychophysical and physiological studies indicate, however, that beyond these early filtering stages, there are stages of 3-D boundary segmentation and surface representation that are used to segregate textures. Chromatic segregation of element-arrangement patterns--as studied by Beck and colleagues--cannot be completely explained by the filtering mechanisms previously employed to account for achromatic segregation. An element arrangement pattern is composed of two types of elements that are arranged differently in different image regions (e.g. vertically on top and diagonally on the bottom). FACADE theory mechanisms that have previously been used to explain data about 3-D vision and figure-ground separation are here used to simulate chromatic texture segregation data, including data with equiluminant elements on dark or light homogeneous backgrounds, or backgrounds composed of vertical and horizontal dark or light stripes, or horizontal notched stripes. These data include the fact that segregation of patterns composed of red and blue squares decreases with increasing luminance of the interspaces. Asymmetric segregation properties under 3-D viewing conditions with the equiluminant elements close or far are also simulated. Two key model properties are a spatial impenetrability property that inhibits boundary grouping across regions with non-collinear texture elements and a boundary-surface consistency property that uses feedback between boundary and surface representations to eliminate spurious boundary groupings and separate figures from their backgrounds.

Color Perception↗

A comparison of the dynamics of simple (Fourier) and complex (non-Fourier) mechanisms in texture segregation.

Models of texture segregation frequently feature two processing mechanisms: simple, linear channels (1st-order, Fourier mechanisms) and complex channels (2nd-order, non-Fourier mechanisms). Using texture patterns designed to segregate primarily as a result of activity in one set of channels or the other, we employed the method of cued response to obtain speed-accuracy tradeoff (SAT) functions measuring the time course of texture segregation processing in simple and complex channels. Here, both simple-channel and complex-channel patterns are composed of Gabor-patch texture elements, thus equating input to simple channels and the first stage of complex channels. Subjects were required to identify the orientation of a rectangular texture-region embedded in a background field of a different texture. SAT functions were obtained by requiring subjects to respond within 200 ms after an auditory cue. We found that: (1) when segregation depended primarily on activity in simple channels, performance was faster and better than when it depended primarily on complex channels; (2) in contrast to a previous study (Sutter, A., & Graham, N. (1995). Investigating simple and complex mechanisms in texture segragation using the speed-accuracy tradeoff method. Vision Research, 35, 2825-2843), simple-channel (Fourier) patterns composed of two textured regions were just as easily segregated as simple-channel patterns in which one of the regions was blank instead of textured; (3) performance with complex-channel patterns composed of diagonally oriented Gabor-patches was considerably worse than performance with complex-channel patterns composed of vertically and/or horizontally oriented Gabor-patches; (4) among simple-channel patterns containing only one region of texture (background-only or rectangle-only), there were minimal differences in performance; and (5) as in previous experiments, there were large individual differences in the segregation of complex-channel (non-Fourier) patterns. All of the above results can be explained within the framework of the simple- and complex-channels model of texture segregation.

Adult↗

Partial segregation of posterior crista and saccular fibers to the nodulus and uvula of the cerebellum in mice, and its development.

The projection of the posterior canal crista and saccular afferents to the cerebellum of embryonic and neonatal mice was investigated using carbocyanine dyes. Anterograde tracing from these two endorgans reveals a partial segregation of these two sets of afferents. The saccule projects predominantly to the uvula, with very minor input to the nodulus. The posterior canal projects mainly to the nodulus and, to a lesser extent, to the uvula. Retrograde tracing from the uvula and nodulus confirms this partial segregation for these two endorgans and extends it to other vestibular endorgans. Uvular injections result in many more labeled fibers in the gravistatic maculae than in the canals' cristae. In contrast, nodular injection reveals many more labeled fibers in the canal cristae than in the gravistatic maculae. This partial segregation may play a role in the information processing in these folia. Our developmental data suggest that the initial segregation at E17 coincides with the formation of the postero-lateral fissure. This embryonic segregation of the primary vestibular mossy fibers to the uvula and nodulus commences long before the maturity of their targets, the granule cells and unipolar brush cells. Thus, the segregation of the primary vestibular projection to the uvula and nodulus does not depend on cues related to the target cells. Rather, the segregation may reflect more global cerebellar patterning mechanisms involving guidance for the vestibular afferent fibers independent of the future target cells.

Aging↗

Application of automated crystallography for transmission electron microscopy in the study of grain-boundary segregation.

Analytical Electron Microscopy (AEM) has brought significant progress in the study of grain-boundary segregation. Using X-ray energy-dispersive spectrometry (XEDS) in the AEM, elemental segregation information can be related to the crystallographic character of the same boundary via conventional Transmission Electron Microscope (TEM) diffraction techniques. While significant efforts have been made to improve XEDS analysis of sub-nanometer segregation layers, the methods for crystallographic characterization of grain boundaries have remained the same for several decades and labor-intensive processes. Recently, a method termed Automated Crystallography for TEM (ACT) was developed, which automates crystallographic characterization of grains under TEM observation. In the present work, we combine ACT and X-ray mapping via EDS in AEM for the study of Sb grain-boundary segregation in a rapidly solidified Cu-0.08 wt % Sb alloy. In contrast with previous reports, a large degree of anisotropy in Sb segregation level between different boundaries is found. ACT results suggest that one of the several grain boundaries observed with no detectable Sb segregation is very close to a Sigma 3 coincidence-site lattice structure. The reason for the observed anisotropy in the present alloy is discussed, based upon McLean's theory of segregation.

Journal Article↗

The segregation of the Escherichia coli origin and terminus of replication.

Escherichia coli chromosome replication forks are tethered to the cell centre. Two opposing models describe how the chromosomes segregate. In the extrusion-capture model, newly replicated DNA is fed bi-directionally from the forks toward the cell poles, forming new chromosomes in each cell half. Starting with the origins, chromosomal regions segregate away from their sisters progressively as they are replicated. The termini segregate last. In the sister chromosome cohesion model, replication produces sister chromosomes that are paired along much of their length. The origins and most other chromosomal regions remain paired until late in the replication cycle, and all segregate together. We use a combination of microscopy and flow cytometry to determine the relationship of origin and terminus segregation to the cell cycle. Origin segregation frequently follows closely after initiation, in strong support of the extrusion-capture model. The spatial disposition of the origin and terminus sequences also fits this model. Terminus segregation occurs extremely late in the cell cycle as the daughter cells separate. As the septum begins to invaginate, the termini of the completed sister chromosomes are transiently held apart at the cell centre, on opposite sides of the cell. This may facilitate the resolution of topological linkages between the chromosomes.

Base Sequence↗

On the mechanism of ooplasmic segregation in single-cell zebrafish embryos.

It has been previously shown that localized elevations of free cytosolic calcium are associated with a morphological contraction in the forming blastodisc and animal hemisphere cortex during ooplasmic segregation in zebrafish zygotes. It was subsequently proposed, in a hypothetical model, that these calcium transients might be linked to the contraction of a cortically located actin microfilament network as a potential driving force for segregation. Here, by labeling single-cell embryos during the major phase of segregation with rhodamine-phalloidin, direct evidence is presented to indicate that the surface contraction was generated by an actin-based cortical network. Furthermore, while zygotes incubated with colchicine underwent normal ooplasmic segregation, those incubated with cytochalasin B did not generate a constriction band or segregate to form a blastodisc. During segregation at the single-cell stage, ooplasm simultaneously moved in two directions: toward the blastodisc within the so-called axial streamers, and toward the vegetal pole in the peripheral ooplasm. The velocities of both axial and peripheral streaming movements are reported. By injection of a fluorescein isothiocyanate (FITC)-labeled 2000 kDa dextran into the peripheral ooplasm it was demonstrated that a portion of it feeds into the bases of the extending streamers, which helps to explain the lack of accumulation of ooplasm at the vegetal pole. These new data were incorporated into the original model to link the bipolar ooplasmic movements with the calcium-modulated, actin-mediated contraction of the animal hemisphere cortex as a means of establishing and driving ooplasmic segregation in zebrafish.

Actins↗

Segregation of hematuria in thin basement membrane disease with haplotypes at the loci for Alport syndrome.

BACKGROUND: Inherited hematuria is common and is usually attributed to thin basement membrane disease (TBMD). The aim of this study was to determine how often hematuria in families with TBMD segregated with haplotypes at the chromosomal loci for autosomal recessive and X-linked Alport syndrome (COL4A3/COL4A4 and COL4A5, respectively). METHODS: The families of 22 individuals with TBMD on renal biopsy and with urinary glomerular red blood cell (RBC) counts of more than 50,000/mL were studied using phase-contrast microscopy of the urine and DNA microsatellite markers. Eighteen families had at least two members with hematuria. RESULTS: Hematuria segregated with or was consistent with segregation at the COL4A3/COL4A4 locus in eight (36%) families (P < 0.05 in 5 of these) and at the COL4A5 locus in four (18%) families (P < 0.05 in 2). The lack of segregation in the other 10 (45%) families may have occurred because of incomplete penetrance of the hematuria, de novo mutations, coincidental hematuria in other family members, or the presence of a novel gene locus. In four different families, three of which had hematuria that segregated with the COL4A3/COL4A4 locus, four family members with the hematuria haplotype had spouses with coincidental hematuria (4 of 29, 14%). However, none of their four offspring who had also inherited the hematuria haplotype had the clinical features of autosomal recessive Alport syndrome. CONCLUSIONS: Hematuria in families with TBMD commonly segregates with the COL4A3/COL4A4 locus and thus results from mutations in the same genes as autosomal recessive Alport syndrome. Sometimes TBMD may be confused with the carrier state for X-linked Alport syndrome. However, nearly half of the families in this study had hematuria that did not segregate with the loci for either autosomal recessive or X-linked Alport syndrome.

Adolescent↗

Gender segregation in childhood: a test of the interaction style theory.

The authors investigated the play/interaction-style theory of gender segregation with a sample of 39 children aged 2 to 5 years (primarily Caucasian). According to this theory, children prefer playmates with styles of play or interaction that are similar to their own. Because such styles are sex differentiated, same-sex playmate preference (i.e., gender segregation) results. The authors observed children during free play to determine preferred playmates and gender segregation level, and they used teacher ratings to derive play/interaction-style scores. The authors used a multiple regression approach to path analysis to analyze effects of sex of participant, participants' play/interaction-style scores, playmates' play/interaction-style scores, and degree of gender segregation to determine their effects on one another. The authors observed significant levels of gender segregation, with highly aggressive or active children displaying less segregation than their peers did. However, gender segregation was not associated with a preference for playmates with similar play or interaction styles.

Child↗

Detection of gene-environment interactions in joint segregation and linkage analysis.

We compare approaches for analysis of gene-environment (G x E) interaction, using segregation and joint segregation and linkage analyses of a quantitative trait. Analyses of triglyceride levels in a single large pedigree demonstrate the two methods and show evidence for a significant interaction (P=.015 when segregation analysis is used; P=.006 when joint analysis is used) between a codominant major gene and body-mass index. Genotype-specific correlation coefficients, between triglyceride levels and body-mass index, estimated from the joint model are rAA=.72, rAa=.49, and raa=. 20. Several simulation studies indicate that joint segregation and linkage analysis leads to less-biased and more-efficient estimates of a G x E-interaction effect, compared with segregation analysis alone. Depending on the heterozygosity of the marker locus and its proximity to the trait locus, we found joint analysis to be as much as 70% more efficient than segregation analysis, for estimation of a G x E-interaction effect. Over a variety of parameter combinations, joint analysis also led to moderate (5%-10%) increases in power to detect the interaction. On the basis of these results, we suggest the use of combined segregation and linkage analysis for improved estimation of G x E-interaction effects when the underlying trait gene is unmeasured.

Body Mass Index↗

Meiotic segregation analysis of RB1 alleles in retinoblastoma pedigrees by use of single-sperm typing.

In hereditary retinoblastoma, different epidemiological studies have indicated a preferential paternal transmission of mutant retinoblastoma alleles to offspring, suggesting the occurrence of a meiotic drive. To investigate this mechanism, we analyzed sperm samples from six individuals from five unrelated families affected with hereditary retinoblastoma. Single-sperm typing techniques were performed for each sample by study of two informative short tandem repeats located either in or close to the retinoblastoma gene (RB1). The segregation probability of mutant RB1 alleles in sperm samples was assessed by use of the SPERMSEG program, which includes experimental parameters, recombination fractions between the markers, and segregation parameters. A total of 2,952 single sperm from the six donors were analyzed. We detected a significant segregation distortion in the data as a whole (P=.0099) and a significant heterogeneity in the segregation rate across donors (.0092). Further analysis shows that this result can be explained by segregation distortion in favor of the normal allele in one donor only and that it does not provide evidence of a significant segregation distortion in the other donors. The segregation distortion favoring the mutant RB1 allele does not seem to occur during spermatogenesis, and, thus, meiotic drive may result either from various mechanisms, including a fertilization advantage or a better mobility in sperm bearing a mutant RB1 gene, or from the existence of a defectively imprinted gene located on the human X chromosome.

Adult↗

QTL analysis of transgressive segregation in an interspecific tomato cross.

Two accessions, representing the species Lycopersicon esculentum (cultivated tomato) and Lycopersicon pennellii (a wild relative), were evaluated for 11 quantitative traits and found to be significantly different for 10 of the traits. Transgressive segregation was observed for eight of the traits in a large interspecific F2 population. When restriction fragment length polymorphism markers were used as probes for the quantitative trait loci (QTL) underlying the traits, 74 significant QTL (LOD > 2) were detected. Thirty-six percent of those QTL had alleles with effects opposite to those predicted by the parental phenotypes. These QTL were directly related to the appearance of transgressive individuals in the F2 for those traits which showed transgressive segregation. However, the same types of QTL (with allelic effects opposite to those predicted by the parents) were also observed for traits that did not display transgressive segregation in the F2. One such trait was dry weight accumulation. When two overdominant QTL (detected in the F2) for this trait were backcrossed into the L. esculentum genetic background, transgressive individuals were recovered and their occurrence was associated with the two QTL demonstrating the potential for transgressive segregation for all characters and implicating overdominance as a second cause of transgressive segregation. Epistasis was not implicated in transgressive segregation in either the F2 or backcross generations. Results from this research not only reveal the basis of wide-cross transgressive segregation, but demonstrate that molecular markers can be used to identify QTL (from wild species) responsible for transgressive phenotypes and to selectively transfer them into crop species. This strategy might be used to improve many traits of economic importance including those for which wild species appear phenotypically inferior to their cultivated counterparts.

Alleles↗

The role of centromere alignment in meiosis I segregation of homologous chromosomes in Saccharomyces cerevisiae.

During meiosis, homologous chromosomes pair and then segregate from each other at the first meiotic division. Homologous centromeres appear to be aligned when chromosomes are paired. The role of centromere alignment in meiotic chromosome segregation was investigated in Saccharomyces cerevisiae diploids that contained one intact copy of chromosome I and one copy bisected into two functional centromere-containing fragments. The centromere on one fragment was aligned with the centromere on the intact chromosome while the centromere on the other fragment was either aligned or misaligned. Fragments containing aligned centromeres segregated efficiently from the intact chromosome, while fragments containing misaligned centromeres segregated much less efficiently from the intact chromosome. Less efficient segregation was correlated with crossing over in the region between the misaligned centromeres. Models that suggest that these crossovers impede proper segregation by preventing either a segregation-promoting chromosome alignment on the meiotic spindle or some physical interaction between homologous centromeres are proposed.

Centromere↗

Nonrandom homolog segregation at meiosis I in Schizosaccharomyces pombe mutants lacking recombination.

Physical connection between homologous chromosomes is normally required for their proper segregation to opposite poles at the first meiotic division (MI). This connection is generally provided by the combination of reciprocal recombination and sister-chromatid cohesion. In the absence of meiotic recombination, homologs are predicted to segregate randomly at MI. Here we demonstrate that in rec12 mutants of the fission yeast Schizosaccharomyces pombe, which are devoid of meiosis-induced recombination, homologs segregate to opposite poles at MI 63% of the time. Residual, Rec12-independent recombination appears insufficient to account for the observed nonrandom homolog segregation. Dyad asci are frequently produced by rec12 mutants. More than half of these dyad asci contain two viable homozygous-diploid spores, the products of a single reductional division. This set of phenotypes is shared by other S. pombe mutants that lack meiotic recombination, suggesting that nonrandom MI segregation and dyad formation are a general feature of meiosis in the absence of recombination and are not peculiar to rec12 mutants. Rec8, a meiosis-specific sister-chromatid cohesin, is required for the segregation phenotypes displayed by rec12 mutants. We propose that S. pombe possesses a system independent of recombination that promotes homolog segregation and discuss possible mechanisms.

Base Sequence↗

Reverse and intermediate segregation of large beads in dry granular media

Mixtures of two types of glass beads have been sheared in a chute flow, in a half-filled rotating drum, and placed in a funnel to form a pile. In the three experimental devices, for small size ratios, there is a segregation of the large beads at the surface of the flowing phase (usual case), but for high size ratios (above about 5) the large beads segregate inside (reverse segregation). Precise measurements show that the segregation drives the large beads to an intermediate level inside the bed. In all devices, there is a continuous evolution of the location of the segregated beads from the surface to deep inside, when increasing the size ratio between the beads. The location of the segregated beads at intermediate levels is well defined both for high size ratios (above 5) and for very small size ratios (about 2), the level being very close to the surface in that case. The reverse and intermediate segregations are masked when using high fractions of large beads in the experiments. Their interpretation involves the high mass of the large particles balancing geometrical effects at a particular intermediate level inside the flowing layer.

Journal Article↗

Probabilistic behavior of DNA segregation in Escherichia coli.

The pattern of segregation of DNA in Escherichia coli B/rK was analyzed by using the Methocel technique for forming chains of cells and the membrane binding elution method. Strain B/rK was shown to have a relatively high degree of nonrandom segregation and was used in a critical experiment to test the proposal that only one DNA strand acts nonrandomly during segregation. Thymidine-labeled cells were bound to a nitrocellulose membrane, and newly dividing cells were eluted from the membrane for six generations. The segregation of DNA in the eluted cells as well as in the cells bound to the membrane was examined by the Methocel technique. No difference in segregation was found between the two populations of cells, a result which indicates that the two strands are equivalent in segregation and that the pattern of segregation is not the result of a permanent binding of any strand to a pole of a cell.

Cell Division↗

Levels of fos, ets2, and myb proto-oncogene RNAs correlate with segregation of chromosome 11 of normal cells and with suppression of tumorigenicity in human cell hybrids.

The tumorigenicity in nude mice of human carcinoma-derived D98AH2 (D98) cells is suppressed when cell hybrids are made by fusing these cells with normal human diploid cells. Selection for hybrids that have segregated chromosomes results in the recovery of tumorigenic segregants. These segregants have all lost at least one copy of chromosome 11 of the diploid cell parent. Earlier we found that the parental D98 cells had detectable levels of mRNA specific for 13 of 21 proto-oncogenes examined. To determine if transregulation of proto-oncogenes by genes of the normal cell occurs in such hybrids, the steady-state levels of mRNA specific to 22 proto-oncogenes in the parental cells were compared with those of nontumorigenic D98 X human diploid hybrids as well as with those of their tumorigenic segregants and with the cells of the resulting tumors. The only chromosome consistently segregated in the latter was chromosome 11 of the diploid cell. fos and ets2 RNA levels and the amount of fos protein were consistently elevated in the segregants compared with amounts in the original hybrids. An unexpected finding was the inverse relationship for myb RNA that was barely detected in the parental D98 cells but was at least 10-fold elevated in hybrids that did not have segregated chromosomes compared with those that did. These patterns were evident in RNAs prepared from both subconfluent and confluent cell cultures. The findings suggest that genes of the normal cell parent can affect proto-oncogene expression. Whether the genes affecting fos, ets2, and myb RNA levels are on chromosome 11 and whether these alterations are causally related to the tumorigenic phenotype of the hybrid remain to be determined.

Animals↗

Chromosome segregation from cell hybrids. II. Do differences in parental cell growth rates and phase times determine direction of loss?

The hypothesis that the direction of chromosome segregation in cell hybrids is determined by the interaction of parent cell cycles, or S-phase times, predicts that the segregant parent will always be the one with the longer cycle, or the longer S phase, and that late replicating chromosomes will be more frequently lost. We have tested this hypothesis by studying cell cycle parameters of mouse, Chinese hamster, and platypus parent cells and by observing chromosome loss and replication patterns in hybrids between them. Two types of hybrids have been studied: mouse-hamster hybrids showed gradual segregation, in one or other direction, of 10-60% chromosomes, while rodent-platypus hybrids (which could be selected under conditions optimal for either parent cell) showed rapid and extreme segregation of platypus chromosomes. We found no correlation between the direction of segregation and the relative lengths of parental cycle times, or phase times, nor between sequence of replication and frequency with which segregant chromosomes are lost. We therefore conclude that the direction and extent of segregation is not directly determined by the interaction of parental cycle or phase times.

Animals↗

Development of the mammalian retinogeniculate pathway: target finding, transient synapses and binocular segregation.

This review is concerned with the development of the mammalian retinogeniculate projection from the perspective of our studies on the hamster and to a lesser extent on the cat. In these, and other mammalian species, axons from the two eyes initially spread throughout the dorsal lateral geniculate nucleus (dLGN) and thus completely overlap. Later they segregate, the axons from each eye coming to occupy discrete, non-overlapping territories within the dLGN. The process of segregation to establish the adult pattern coincides with the death of retinal ganglion cells projecting to inappropriate areas of the dLGN and with the loss, by degeneration or retraction, of the axons and/or axonal branches initially located within inappropriate territory of the dLGN. These events occur in the early postnatal period in hamsters, before the eyes have opened, and in cats and monkeys they occur entirely during embryonic life: thus, they do not depend on the onset of normal visual function. If one eye is removed before segregation has begun, the terminal fields of the crossed and uncrossed axons from the remaining eye do not segregate, suggesting that segregation in normal development may depend on some form of interaction between retinal ganglion cells from the two eyes. Attractive and/or repulsive influences exerted by the dLGN on retinogeniculate axons may also be involved in the formation of eye-specific territories. Experimental ultrastructural studies in hamster and cat show that the overlap phase is associated with the formation, by inappropriately located axons, of transient synapses similar to those made by retinogeniculate axons in appropriate parts of the dLGN. In the cat, the transient synapses are made by the axon trunk and by side branches of retinogeniculate axons with terminal arbors in appropriate parts of the nucleus; the transient synapses disappear as the side branches are shed or retracted during the segregation period. Because of good evidence that electrical activity of the retinogeniculate axons may be involved in binocular segregation of inputs, we suggest that the formation and elimination of transient synapses play a significant role in the development of the orderly retinogeniculate projections.

Aging↗