Patch Clustering Research Articles

Maximizing intake under challenging foraging conditions at two spatial scales in Soay sheep

We examined how decision-making processes at small spatial scales can account for individual movements and food intake at larger scales. Our experiment used two currencies that might affect animal decisions: food reward at the patch scale and spatial distribution of the patches. We used Soay sheep, Ovis aries, as the model species in conjunction with an artificial arena that allowed us to modify food-handling conditions and the distribution of the food patches to record variation in food intake of both sexes. The main variable that affected food intake was how efficient the animals were at avoiding patches that were depleted in previous visits. Intake was also affected by mouth size and the spatial distribution of the feeding stations. Sheep were more efficient at exploiting clustered patches with a high energy reward than a matrix of homogeneously distributed patches of lower energy reward. Females' intake was higher than males'; we discuss this in relation to different foraging strategies related to sexual differences in fibre digestibility and the distribution of food resources of different quality that both sexes might use in field conditions. The results indicated that variation in food-handling conditions at the patch level affects food intake at larger scales, providing support for the hypothesis that animal movement related to grazing activity is a hierarchical process, scaling up from bite size to large spatial scales. However, spatial memory is important for maximizing the use of patchily distributed food.

Seagrass as pasture for seacows: Landscape-level dugong habitat evaluation

A 24 km 2 seagrass meadow in Hervey Bay, Queensland, Australia, was confirmed as important dugong habitat by a satellite tracking study. Marine videography, near-infrared spectroscopy (NIRS) and geographic information systems (GIS) were used to survey, analyse and map seagrass species composition, nutrient profile and patch structure at high resolution (200 m). Five species of seagrass covered 91% of the total habitat area. The total above and below-ground seagrass biomass was estimated to be 222.7 ± 19.6 t dry-weight. Halodule uninervis dominated the pasture (81.8%, 162.2 t), followed by Halophila ovalis (35.3%, 16.5 t), Zostera capricorni (15.9%, 22.2 t), Halophila spinulosa (14.5%, 21.9 t), and traces of Halodule pinifolia. Because seagrass distributions overlapped, their combined percentage totalled >100% of the survey area. The seagrass formed a continuous meadow of varying density. Abiotic variables explained relatively little of the spatial patterns in the seagrass. For all seagrass species, the above-ground component (shoots and leaves) possessed greater total nitrogen than the below-ground component (roots and rhizomes), which possessed greater total starch. Because of the relatively low intraspecific variation in nutrient composition, nutrients were concentrated according to seagrass biomass density. Halodule uninervis was the most nutritious seagrass species because of its superior whole-plant nitrogen (1.28 ± 0.05% DW) and starch (6.42 ± 0.50 DW %) content. Halodule uninervis formed large, clustered patches of dense biomass across the pasture and thus nitrogen and starch were concentrated where H. uninervis was prevalent. This seagrass meadow appears to be utilised well below its potential dugong carrying capacity. The survey and analytical techniques used enabled rapid, economical and accurate quantification and characterisation of seagrass habitat at scales relevant to a large forager.

PI(4,5)P2-dependent microdomain assemblies capture microtubules to promote and control leading edge motility

The lipid second messenger PI(4,5)P2 modulates actin dynamics, and its local accumulation at plasmalemmal microdomains (rafts) might mediate regulation of protrusive motility. However, how PI(4,5)P2-rich rafts regulate surface motility is not well understood. Here, we show that upon signals promoting cell surface motility, PI(4,5)P2 directs the assembly of dynamic raft-rich plasmalemmal patches, which promote and sustain protrusive motility. The accumulation of PI(4,5)P2 at rafts, together with Cdc42, promotes patch assembly through N-WASP. The patches exhibit locally regulated PI(4,5)P2 turnover and reduced diffusion-mediated exchange with their environment. Patches capture microtubules (MTs) through patch IQGAP1, to stabilize MTs at the leading edge. Captured MTs in turn deliver PKA to patches to promote patch clustering through further PI(4,5)P2 accumulation in response to cAMP. Patch clustering restricts, spatially confines, and polarizes protrusive motility. Thus, PI(4,5)P2-dependent raft-rich patches enhance local signaling for motility, and their assembly into clusters is regulated through captured MTs and PKA, coupling local regulation of motility to cell polarity, and organization.

Spatial control of actin-based motility through plasmalemmal PtdIns(4,5)P2-rich raft assemblies.

The interactions of cells with their environment involve regulated actin-based motility at defined positions along the cell surface. Sphingolipid- and cholesterol-dependent microdomains (rafts) order proteins at biological membranes, and have been implicated in most signalling processes at the cell surface. Many membrane-bound components that regulate actin cytoskeleton dynamics and cell-surface motility associate with PtdIns(4,5)P(2)-rich lipid rafts. Although raft integrity is not required for substrate-directed cell spreading, or to initiate signalling for motility, it is a prerequisite for sustained and organized motility. Plasmalemmal rafts redistribute rapidly in response to signals, triggering motility. This process involves the removal of rafts from sites that are not interacting with the substrate, apparently through endocytosis, and a local accumulation at sites of integrin-mediated substrate interactions. PtdIns(4,5)P(2)-rich lipid rafts can assemble into patches in a process depending on PtdIns(4,5)P(2), Cdc42 (cell-division control 42), N-WASP (neural Wiskott-Aldrich syndrome protein) and actin cytoskeleton dynamics. The raft patches are sites of signal-induced actin assembly, and their accumulation locally promotes sustained motility. The patches capture microtubules, which promote patch clustering through PKA (protein kinase A), to steer motility. Raft accumulation at the cell surface, and its coupling to motility are influenced greatly by the expression of intrinsic raft-associated components that associate with the cytosolic leaflet of lipid rafts. Among them, GAP43 (growth-associated protein 43)-like proteins interact with PtdIns(4,5)P(2) in a Ca(2+)/calmodulin and PKC (protein kinase C)-regulated manner, and function as intrinsic determinants of motility and anatomical plasticity. Plasmalemmal PtdIns(4,5)P(2)-rich raft assemblies thus provide powerful organizational principles for tight spatial and temporal control of signalling in motility.

Apical localization of actin patches and vacuolar dynamics in Ashbya gossypii depend on the WASP homolog Wal1p.

Analysis of the Ashbya gossypii Wiskott-Aldrich syndrome-like gene AgWAL1 indicates that it is required for the maintenance of polarized hyphal growth. Growth and organelle dynamics of the wild type and of wal1 and other mutant strains were monitored by in vivo (fluorescence) time-lapse microscopy. Loss of WAL1 led to slow growth and defects in polarized growth that produced swellings in subapical regions, whereas formation of hyphal tips and dichotomous tip branching occurred as in the wild-type. Few actin cables in Agwal1 cells were found to insert into the hyphal tip, but specific clustering of cortical actin patches was observed in subapical regions of hyphal tips instead of at the hyphal apex. Distribution and movement of vacuoles was observed in vivo using FM4-64. In the wild type and in the slowly growing mutant strains bem2 and cla4, which lack a Rho-GTPase-activating protein and a PAK kinase, respectively, early endosomes appeared in the hyphal tip, whereas very few early endosomes and small vacuoles were found in the wal1 mutant hyphal tips, thus linking the cortical patch defect of wal1 hyphae with the distribution of endosomes. Vivid movement of vacuoles seen in the wild type and in the bem2 mutant in subapical regions was largely reduced in the wal1 and cla4 mutants. The tubular structure of mitochondria (as visualized by DIOC6 in vivo) was similar in the wild type and the wal1 mutant, although wal1 mitochondria appeared to be larger. Interestingly, mitochondria were found to insert into the hyphal tips in both strains. Our results indicate a function for Wal1p in filamentous fungi in coordinating actin patch distribution with polarized hyphal tip growth.

On optimal choices in increase of patch area and reduction of interpatch distance for metapopulation persistence

Metapopulation theory teaches that the viability of metapopulations may be enlarged by decreasing the probability of extinction of local populations, or by increasing the colonization probability of empty habitat patches. In a metapopulation model study it has recently been found that reducing the extinction probability of the least extinction-prone patch and increasing the colonization probability between the two least extinction-prone patches are the best options to prolong the expected lifetime of a metapopulation. In this article I examine with a more detailed model whether this translates into enlarging the largest patch and reducing the interpatch distance between the largest patches. Using two measures of metapopulation persistence (longevity and resilience) I found, firstly, that the largest patch should generally be enlarged if the choice is to enlarge a patch by a certain percentage (relative change) and that the smallest patch should generally be enlarged if the choice is to enlarge a patch by a fixed amount of area (absolute change), and secondly that indeed one should reduce the interpatch distance between the two largest patches, if the choice is among all pairs of patches. The strength of these rules of thumb (particularly the second part of the first rule) depends on the parameter values, particularly the amount of clustering of patches, the mean dispersal distance and the number of dispersers. Also, the rules of thumb are less pronounced when resilience is chosen as a measure of metapopulation persistence than when longevity is chosen.

Spatial Pattern Analysis of Sharka Disease (Plum pox virus Strain M) in Peach Orchards of Southern France

ABSTRACT The spatial pattern of Sharka disease, caused by Plum pox virus (PPV) strain M, was investigated in 18 peach plots located in two areas of southern France. PPV infections were monitored visually for each individual tree during one to three consecutive years. Point pattern and correlation-type approaches were undertaken using the binary data directly or after parsing them in contiguous quadrats of 4, 9, and 16 trees. Ordinary runs generally revealed a low but variable proportion of rows with adjacent symptomatic trees. Aggregation of disease incidence was indicated by the theta parameter of the beta-binomial distribution and related indices in 15 of the 18 plots tested for at least one assessment date of each. When aggregation was detected, it was indicated at all quadrat sizes and tended to be a function of disease incidence, as shown by the binary form of Taylor's power law. Spatial analysis by distance indices (SADIE) showed a nonrandom arrangement of quadrats with infected trees in 14 plots. The detection of patch clusters enclosing quadrats with above-average density of symptomatic trees, ellipsoidal in shape and generally extending from 4 to 14 trees within rows and from 4 to 10 trees perpendicular to the rows, could be interpreted as local areas of influence of PPV spread. Spatial patterns at the plot scale were often characterized by the occurrence of several clusters of infected trees located up to 90 m apart in the direction of the rows. When several time assessments were available, increasing clustering over time was generally evidenced by stronger values of the clustering index and by increasing patch cluster size. The combination of the different approaches revealed a wide range of spatial patterns of PPV-M, from no aggregation to high aggregation of symptomatic trees at all spatial scales investigated. Such patterns suggested that aphid transmission to neighboring trees occurred frequently but was not systematic. The mechanism of primary virus introduction, the age and structure of the orchards when infected, and the diversity of vector species probably had a strong influence on the secondary spread of the disease. This study provides a more complete understanding of PPV-M patterns which could help to improve targeting of removal of PPV-infected trees for more effective disease control.

Are pre-Columbian conditions relevant baselines for managed forests in the northeastern United States?

Populations of a number of taxa associated with shrublands, early-successional forests, and other disturbance-generated habitats (collectively referred to as thickets) are declining in the northeastern United States. To assure that species dependent on thicket habitats persist, intervention is warranted. However, conservationists concerned with the status of thicket-dependent species are confronted with two important questions. How much habitat is needed? And how should these habitats be distributed? Natural disturbance regimes have been recommended as a baseline that managers should consider while providing thicket habitats. Within the Northeast, historic disturbance regimes varied substantially among forest types. Coastal regions were characterized by extensive barrens where regular and often times large-scale disturbances that resulted in >15% of the area being covered by regenerating forest stands. Among inland forests, natural disturbances were usually small and resulted in seedling-sapling stands and beaver ( Castor canadensis) impoundments covering <6% of the area. Under these conditions, thicket-affiliated species were probably distributed in small, disjunct populations that shifted in space and time. Current efforts to maintain thicket habitats must deal with a range of current land-uses and a legacy of historic uses. Additionally, the effectiveness of management protocols that mimic natural disturbances is limited among many forests. Increasing ownership parcelization, a relatively young forest, and landscape fragmentation substantially reduce the practicality and suitability of small-scale disturbances for generating thicket habitats. Large, clustered patches may be more practical and beneficial, especially in urbanized landscapes. In rural areas, silvicultural manipulations should be applied on a “sliding scale” relative to forest age. Timber harvests that emulate the range of variability of natural disturbances may become appropriate in these areas as forest stands mature. Addressing the needs of thicket-dependent species in the northeastern United States will require creativity, a willingness to explore a variety of solutions, and public support.

Biogeochemical Changes Accompanying Woody Plant Encroachment in a Subtropical Savanna

Ecosystem properties of surficial (0–10 cm) soils in remnant herbaceous patches were compared to those of contrasting woody plant patch types (upland discrete cluster, upland grove, and lowland woodland) where shifting land cover is known to have occurred over the past 50–77 yr. The purpose of this study was to evaluate and quantify the biogeochemical consequences and subsequent developmental rates of woody plant formation on sites formerly dominated by grasses. Clay and water content of woodland soil patches was higher than that of soils associated with upland discrete cluster and grove patches. Even so, lowland woody patches were generally comparable to upland grove and discrete shrub cluster patches with respect to soil organic carbon (SOC), soil N, the ratio of annual N mineralization:total N, annual litterfall, and root biomass. The fact that finer soil texture, enhanced soil moisture, and the more advanced age of lowland woody patches did not translate into greater accumulations of SOC and N relative to upland grove and discrete cluster patches suggests that C and N losses might be higher in recently developed lowland woodland communities. Fluctuations in monthly root biomass standing crop (0–10 cm) far exceeded annual foliar litterfall in upland and lowland woody patch types, suggesting that belowground inputs of organic matter may drive changes in soil physical and chemical properties that occur subsequent to woody plant establishment. The estimated annual mean rates of soil C accretion in the “islands of fertility” that developed subsequent to tree/shrub encroachment were variable and ranged from 8 to 23 g/m2 (in groves and discrete clusters, respectively); N accretion ranged from 0.9 to 2.0 g/m2 (in groves and discrete clusters, respectively), even though mean annual N mineralization rates were three- to fivefold greater than those measured in remnant herbaceous patches. Woody plant proliferation in grasslands and savannas in recent history has been widely reported around the world. The causes for this shift in vegetation are controversial and center around changes in livestock grazing, fire, climate, and atmospheric CO2. Our data, which are conservative in that they examine only the upper 10 cm of the soil profile, indicate that the rate and extent of soil C and N accumulation associated with this phenomenon can be rapid, substantial, and accompanied by increased N turnover. This geographically extensive vegetation change thus has important implications for understanding how the global carbon and nitrogen cycles may have been altered since Anglo-European settlement of arid and semiarid regions.

BIOGEOCHEMICAL CHANGES ACCOMPANYING WOODY PLANT ENCROACHMENT IN A SUBTROPICAL SAVANNA

Ecosystem properties of surficial (0–10 cm) soils in remnant herbaceous patches were compared to those of contrasting woody plant patch types (upland discrete cluster, upland grove, and lowland woodland) where shifting land cover is known to have occurred over the past 50–77 yr. The purpose of this study was to evaluate and quantify the biogeochemical consequences and subsequent developmental rates of woody plant formation on sites formerly dominated by grasses. Clay and water content of woodland soil patches was higher than that of soils associated with upland discrete cluster and grove patches. Even so, lowland woody patches were generally comparable to upland grove and discrete shrub cluster patches with respect to soil organic carbon (SOC), soil N, the ratio of annual N mineralization:total N, annual litterfall, and root biomass. The fact that finer soil texture, enhanced soil moisture, and the more advanced age of lowland woody patches did not translate into greater accumulations of SOC and N relative to upland grove and discrete cluster patches suggests that C and N losses might be higher in recently developed lowland woodland communities. Fluctuations in monthly root biomass standing crop (0–10 cm) far exceeded annual foliar litterfall in upland and lowland woody patch types, suggesting that belowground inputs of organic matter may drive changes in soil physical and chemical properties that occur subsequent to woody plant establishment. The estimated annual mean rates of soil C accretion in the “islands of fertility” that developed subsequent to tree/shrub encroachment were variable and ranged from 8 to 23 g/m2 (in groves and discrete clusters, respectively); N accretion ranged from 0.9 to 2.0 g/m2 (in groves and discrete clusters, respectively), even though mean annual N mineralization rates were three- to fivefold greater than those measured in remnant herbaceous patches. Woody plant proliferation in grasslands and savannas in recent history has been widely reported around the world. The causes for this shift in vegetation are controversial and center around changes in livestock grazing, fire, climate, and atmospheric CO2. Our data, which are conservative in that they examine only the upper 10 cm of the soil profile, indicate that the rate and extent of soil C and N accumulation associated with this phenomenon can be rapid, substantial, and accompanied by increased N turnover. This geographically extensive vegetation change thus has important implications for understanding how the global carbon and nitrogen cycles may have been altered since Anglo-European settlement of arid and semiarid regions.

The role of DOM sorption to mineral surfaces in the preservation of organic matter in soils

Sorption of dissolved organic matter (DOM) is considered to be a major process in the preservation of organic matter (OM) in marine sediments. Evidence for this hypothesis includes the close relationship between sediment surface area (SA) and organic carbon (OC) concentrations and the strongly reduced biological degradability after DOM has sorbed to mineral surfaces. The aim of this study was to discuss the possibility of a similar process in the soil environment. We accomplished this by gathering information from the literature, and by an evaluation of our own studies on DOM sorption and accumulation of OM in soil. We found that in soil a close association of OM with the mineral matrix exists. Both the concentration of soil OM associated with the mineral matrix, and the sorption of DOM are related to reactive mineral phases such as Al and Fe oxyhydroxides. Sorption of DOM derived from the oxidative decomposition of lignocellulose to Al and Fe oxyhydroxides involves strong complexation bondings between surface metals and acidic organic ligands, particularly with those associated with aromatic structures. The strength of the sorption relates to the surface area but more importantly to the surface properties of the sorbing mineral phase. The sorption of a large part of DOM is hardly reversible under conditions similar to those during sorption (hysteresis). Because sorption of the more labile polysaccharide-derived DOM on mineral surfaces is weaker, adsorptive and desorptive processes strongly favour the accumulation of the more recalcitrant lignin-derived DOM. In addition, we found the soil OM in an alluvial B horizon and in the clay fraction of a topsoil strongly resembling lignin-derived DOM from the overlying forest floors. Hence, it seems likely that sorption of DOM contributes considerably to the accumulation and preservation of OM in soil. However, this does not result in a significant relationship between OC concentration and SA. Reasons for that finding may be the ”masking” of mineral surfaces by adsorbed OM, the clustering of OM patches at highly reactive sites of metal hydroxides, and/or the absence of a relationship between SA and the concentration of surface-active Fe and Al oxyhydroxides in some soil types. Overall, we conclude that sorptive preservation of OM in soil is affected by the chemical structure of the sorbing DOM and the surface properties of the mineral matrix. Localisation and conformation of sorbed OM remains unclear and therefore should be subject of further research.

Strengths abd weaknesses of the reef guild concepts and quantitative data: Application to the upper Capitan-massive community (Permian), Guadalupe Mountains, New Mexico-Texas

Analyses of large acatate sheet tracings, close-up photos and 105 sub-horizontal quadrat surfaces at four localities near the base of the Guadalupe Mountains Escarpment indicate that the biotic framework of the upper Capitan reef was built by about 35 species: one codiacean (Eugonophyllum sp.), 17 calcisponges, 9 bryozoans, one richthofenid brachipod, some crinoid (known only from columns), 4 Problematica and microbes. This widespread fossil community included members of the Constructor, Baffler and Binder Guilds. A re-evaluation of the Guild Concept (Fagerstrom, 1987, 1991) highlights the validity of the functional roles of the Constructor and Binder Guilds for reef construction. Members of the Baffler Guild, however, need to be revised and an interpretation of microbial micrite and cryptic biota remains controversial. Open surface phylloid algal and cryptic sponge-bryozoan dominated sub-communities were of only local importance. The upper Capitanmassive differs from its Permian conterparts in the low diversity and areal cover of the frame-building biota, low micrite content and abundant micro-frameworks, i.e, intergrown small sponges, Problematica and syndepositional cements (botryoidal and isopachous, fibrous calcite). Quantitative areal cover data were assessed at various scales. Large acetate sheets generally have low coverage of macro-biota (5.4%). By contrast, analysis of small areas of local high areal cover (selected acetate sheet quadrats, subvertical photographs, and quadrat samples: 15–21%) provide detailed insights into clustered patches forming the inital reef framework. Both data sets provide useful clues for an integrated approach to framework assessment. Mean acetate sheet data are limited by their somewhat generalized pattern, while small investigation areas may overemphasize local variation. Erect and pendant sponges with solitary, sub-cylindrical and multi-branche/clonal forms, were the predominant initial frame-builders in both open surface and cryptic habitats. Selective larval recruitment of erects sponges to firm substrates produced continous upward accretion of the initial framework. On open surfaces and in pores formed by tabular sponges and fenestrate bryozoans, erect and pendant sponges were supported in their hydrodynamically unstable growth position by encrusters, chieflyArchaeolithoporelle hidensis, Shamovella obscura, an unnamed tubular organism, and microbes. Subsequent growth ofArchaeolithoporella hidensis, microbial crusts and syndepositional cements on the outer walls of live sponges would have impeded ambient water circulation and may have led to ‘creeping sponge death by suffocation’ or complete encrustation after death. Filling of pores in the initial and encrusted reef framework by internal sediment (packstone-grainstone; derived from the framework and the back-reef shelf/platform) and voluminous syndepositional marine-phreatic cements completed the framebuilding process.

Short-term response of wildlife to clear-cutting in Quebec boreal forest: multiscale effects and management implications

We studied the short-term response of wildlife to clear-cutting in four blocks (52-114 km2) that were logged in 100- to 250-ha clustered patches. Surveys were conducted, 2 years before and 2 years after logging, to determine the relative abundance of 12 wildlife species, and telemetry data were also gathered on four species. Small mammals, species with the smallest home ranges ([Formula: see text]1 ha), either remained in the clearcut patches or had replacement habitat in the buffer strips. Most species with home ranges up to 25 ha (spruce grouse, Falcipennis canadensis Linné; snowshoe hare, Lepus americanus Erxleben) were excluded from clearcut patches. Species with home ranges [Formula: see text]5 km2 (marten, Martes americana Turton; moose, Alces alces Linné) remained in some residual forest patches scattered throughout clearcuts and in the adjacent uncut forest. In their home range, these two species avoided clearcut patches where the shrub layer and coniferous regeneration were scattered. Because many wildlife species depend on residual forest, the important issue is not the size of clearcut patches but the extent and configuration of the remaining forest. Instead of a clustered distribution of clearcut patches, we propose two harvest scenarios more compatible with integrated wildlife-forest management objectives on a local scale.

Neural (N)‐cadherin at developing thalamocortical synapses provides an adhesion mechanism for the formation of somatopically organized connections

Thalamic projections from the ventrobasal (VB) nucleus to rodent somatosensory cortex develop highly ordered terminations that form discrete, clustered patches localized to layer IV cellular aggregates that are termed “barrels.” The molecular signaling and adhesion events that occur at the synapse as barrel-clustered thalamic connections form are unknown. Here, we show that neural (N)-cadherin, a membrane glycoprotein mediating strong homophilic adhesion, is concentrated at the developing thalamocortical synaptic junctional complex and demarcates these synaptic junctions as they form their characteristic barrel clusters during the first postnatal week. Furthermore, experimentally altering the distribution of thalamocortical axon terminals by peripheral manipulation leads to an identically altered N-cadherin distribution pattern, which is significant in establishing that N-cadherin does not define region-specific patterns of synapse distribution proactively but, rather, conforms to patterning imposed by thalamic axons through instructional cues conveyed through several synaptic relays. At postnatal day 9, levels of N-cadherin expression rapidly decrease, leading to loss of N-cadherin labeling of the barrels and, at adulthood, elimination from VB thalamocortical synapses. However, αN- and β-catenin, which are critical binding partners of the classic cadherins, persist at the adult synapse, suggesting the presence of another classic cadherin as the thalamocortical synapse matures. This is the first evidence linking a synapse adhesion molecule with the establishment of patterned thalamocortical synapse distribution, suggesting strongly that N-cadherin performs a critical role in this process by adhering presynaptic and postsynaptic membranes as ingrowing thalamic axon terminals and postsynaptic thalamorecipient sites link and stabilize into mature synaptic junctional complexes distributed with precise topographic order. It is speculated that the developmental redistribution of N-cadherin may reflect dynamic regulation of synaptic membrane adhesion, which, in turn, might modulate plasticity of thalamocortical synaptic function. J. Comp. Neurol. 407:453–471, 1999. © 1999 Wiley-Liss, Inc.

Neural (N)-cadherin at developing thalamocortical synapses provides an adhesion mechanism for the formation of somatopically organized connections

Thalamic projections from the ventrobasal (VB) nucleus to rodent somatosensory cortex develop highly ordered terminations that form discrete, clustered patches localized to layer IV cellular aggregates that are termed "barrels." The molecular signaling and adhesion events that occur at the synapse as barrel-clustered thalamic connections form are unknown. Here, we show that neural (N)-cadherin, a membrane glycoprotein mediating strong homophilic adhesion, is concentrated at the developing thalamocortical synaptic junctional complex and demarcates these synaptic junctions as they form their characteristic barrel clusters during the first postnatal week. Furthermore, experimentally altering the distribution of thalamocortical axon terminals by peripheral manipulation leads to an identically altered N-cadherin distribution pattern, which is significant in establishing that N-cadherin does not define region-specific patterns of synapse distribution proactively but, rather, conforms to patterning imposed by thalamic axons through instructional cues conveyed through several synaptic relays. At postnatal day 9, levels of N-cadherin expression rapidly decrease, leading to loss of N-cadherin labeling of the barrels and, at adulthood, elimination from VB thalamocortical synapses. However, alphaN- and beta-catenin, which are critical binding partners of the classic cadherins, persist at the adult synapse, suggesting the presence of another classic cadherin as the thalamocortical synapse matures. This is the first evidence linking a synapse adhesion molecule with the establishment of patterned thalamocortical synapse distribution, suggesting strongly that N-cadherin performs a critical role in this process by adhering presynaptic and postsynaptic membranes as ingrowing thalamic axon terminals and postsynaptic thalamorecipient sites link and stabilize into mature synaptic junctional complexes distributed with precise topographic order. It is speculated that the developmental redistribution of N-cadherin may reflect dynamic regulation of synaptic membrane adhesion, which, in turn, might modulate plasticity of thalamocortical synaptic function.

Scaling properties in landscape patterns: New Zealand experience

In this paper we present a case study of spatial structure in landscape patterns for the North and South Islands of New Zealand. The aim was to characterise quantitatively landscape heterogeneity and investigate its possible scaling properties. The study examines spatial heterogeneity, in particular patchiness, at a range of spatial scales, to help build understanding on the effects of landscape heterogeneity on water movement in particular, and landscape ecology in general. We used spatial information on various landscape properties (soils, hydrogeology, vegetation, topography) generated from the New Zealand Land Resource Inventory. To analyse this data set we applied various methods of fractal analyses following the hypothesis that patchiness in selected landscape properties demonstrates fractal scaling behaviour at two structural levels: (1) individual patches; and (2) mosaics (sets) of patches. Individual patches revealed scaling behaviour for both patch shape and boundary. We found self-affinity in patch shape with Hurst exponent H from 0.75 to 0.95. We also showed that patch boundaries in most cases were self-similar and in a few cases of large patches were self-affine. The degree of self-affinity was lower for finer patches. Similarly, when patch scale decreases the orientation of patches tends to be uniformly distributed, though patch orientation on average is clearly correlated with broad scale geological structures. These results reflect a tendency to isotropic behaviour of individual patches from broad to finer scales. Mosaics of patches also revealed fractal scaling in the total patch boundaries, patch centers of mass, and in patch area distribution. All these reflect a special organisation in patchiness represented in fractal patch clustering. General relationships which interconnect fractal scaling exponents were derived and tested. These relationships show how scaling properties of individual patches affect those for mosaics of patches and vice-versa. To explain similarity in scaling behaviour in patchiness of different types we suggest that the Self-Organised Criticality concept should be used. Also, potential applications of our results in landscape ecology are discussed, especially in relation to improved neutral landscape models.

The effects of experimental habitat destruction and patch isolation on space use and fitness parameters in female root vole Microtus oeconomus.

The aim of this study was to test whether experimentally induced destruction and fragmentation of habitat and varying degrees of patch isolation applied to 11 experimental populations of root voles affected space use and fitness parameters in radio-tracked females. It was predicted that the habitat destruction would disrupt the spatiosocial relations between breeding individuals, which, in turn, would cause lowered survival and reproductive success. Different degrees of patch isolation were expected to affect the rate of risky and energetically costly interpatch movements. The experimental habitat destruction comprised the removal of more than half of the habitat area (meadow vegetation) in each of seven populations that had been established in a large continous habitat block. This treatment yielded six small, habitat patches arranged in two clusters with long and short interfragment distances, respectively. Measures of individual space use, survival rates (predation and unknown causes) and litter production were contrasted with equivalent measures obtained from four control populations. The control populations had been established in permanently fragmented plots with the same fragment configuration as the post-destruction treatment populations. The effect of different interpatch distance was tested by comparing movement rates, space use and demographic parameters between the two types of patch cluster. Home ranges were larger in the continuous predestruction populations compared to the permanently fragmented control populations. There was a large decrease in home range area of individual females as a result of habitat destruction, and they became temporally smaller than in the control plots. The degree of overlap only increased on home range core areas. The degree of space sharing between matrilineal related females, a main characteristic of Microtus social organization, was higher in the permanently fragmented populations (controls) than in the treatment populations. This difference was also maintained after habitat destruction. Thus, the basic spatio-social organization of reproductive females when first established, seems to be very resistant to habitat destruction. There were no effects of habitat destruction on female fitness parameters when tested for either at the cluster or the population level. There were less between-patch movements when interpatch distances were large than when they were small, irrespective of the fragmentation history of the population. Predation by avian predators, the main mortality cause, was highest in fragment clusters with long interpatch distances, probably because long distance movements between fragments increased the predation risk. There was a large intrapopulation variation with respect to how severely individuals were affected by the habitat destruction. The farther a female had to relocate her home range because of habitat destruction, the higher was the predation risk. Thus, this study predicts that the most influential aspect of habitat fragmentation on demography may be that causing long interpatch movements.

Chapter 10 The role of nitric oxide in development of the patch—cluster system and retinocollicular pathways in the rodent superior colliculus

Nitric oxide (NO) has been implicated as a retrograde signal in the process of refining axonal pathways during brain development. To determine some of the factors involved in this process, we have used two model pathway systems in the rat and mouse superior colliculus (SC). The first, the patch-cluster system, consists of clusters of neurons in the intermediate gray layer (igl) which transiently express NO during development and which receive input from a cholinergic pathway from the parabrachial brainstem as well as from other pathways containing different transmitters. The second system, the retinocollicular pathway, consists of glutamatergic fibers that project to the superficial gray layer. We have used both nitric oxide synthase inhibition (nw-nitro-L-arginine, NoArg) and single (nNOS) and double (nNOS and eNOS) gene knockout mice to examine the effect that reduction in NOS has upon the development of these two systems. The onset of NOS expression in rat, as revealed by nicotinamide adenine dinucleotide phosphate diaphorase (NADPH-d) labeling, occurred in igl cells as early as postnatal day P5, with clusters being well-established by P14. Cholinergic fibers were first visible at P10 and formed obvious patches and tiers by P14. Intraperitoneal injections of NoArg from P1-P22 had no effect upon the development of these cholinergic patches. The pathway also developed normally in both single and double-knockout mice. In contrast, the ipsilateral retinocollicular pathway was altered in the double, but not in the single knockout mouse. This pathway is exuberant during the first week of life, being distributed across much of the mediolateral axis of the rostral SC. By P8-P15, this pathway has retracted to the most mediorostral SC. This refinement was delayed substantially in the double NOS gene knockout mouse. Ipsilateral fibers were found within 3-5 separate medio-lateral patches within the rostral 600 microns of SC at P15, and patches of abnormal size and extent were also seen at P18. We conclude from these results that NO plays a role in pathway development in the rodent SC, but only in glutamatergic pathways and only when both endothelial and neuronal forms of NOS have been deleted. The mechanism of this effect must involve pathway elimination in situations where there is non-correlated electrical activity. It is likely that NO promotes fiber retraction rather than fiber stabilization in these developing nerve fibers.

Dystrophin-glycoprotein complex and vinculin-talin-integrin system in human adult cardiac muscle

Costameres were identified, for the first time, in skeletal and cardiac muscle, as regions associated with the sarcolemma, consisting of densely clustered patches of vinculin; they have many characteristics common to the cell-extracellular matrix-type of adherens junctions. Costameres are considered 'proteic machinery' and they appear to comprise two protein complexes, the dystrophin-glycoprotein complex (DGC) and the vinculin-talin-integrin system. In comparison to skeletal muscle, few studies have focused on cardiac muscle regarding these two complexes, and study is generally relative to dystrophin or to cardiac diseases, such as cardiomyopathies. However, insufficient data are available on these proteins in healthy human cardiomyocytes. For this reason, we performed an immunohistochemical study using human cardiac muscle fibers, in order to define the real distribution and the spatial relationship between the proteins in these two complexes. Our data showed a real costameric distribution of DGC and of the vinculin-talin-integrin system; all tested proteins were present in T-tubule and in intercalated disks. Moreover, our data demonstrated that all tested proteins of DGC colocalized with each other, as all tested components of the vinculin-talin-integrin system, and that all tested proteins of DGC colocalized with all tested proteins of the vinculin-talin-integrin system. Finally, all tested proteins of the two complexes were localized in the region of the sarcolemma over the I band, in 100% of our observations. The present study, for the first time, analyzed the majority of proteins of DGC and of the vinculin-talin-integrin system in cardiac muscle fibers, and it confirmed that DGC and the vinculin-talin-integrin system have a role in the transduction of mechanical force to the extracellular matrix. Finally it attributed a key role in the regulation of action potential duration to cardiac myocytes.

Effect of dimerization of the d-glucose analogue of muramyl dipeptide on stimulation of macrophage-like cells

N-Acetylmuramyl- l-alanyl- d-isoglutamine (MDP) is the minimum required structure responsible for the immunoadjuvant activity of the bacterial cell wall. The d-glucose analogue of MDP (GADP) was reported to show a higher immunoadjuvant activity than MDP itself. Although the mechanism of activation by MDP and the existence of receptor against MDP are not clear, the patch formation and cluster formation of receptors are important steps on the signal transduction by such bioactive molecules. It is expected that the cluster effect such as antennary oligosaccharides reported by Lee et al. increased the affinity of ligand against receptor and accelerated the patch formation and cluster formation of receptors. In order to discuss the effect of multivalent-ligand formation of GADP on the activation of immunocompetent cells in more detail, we have synthesized GADP dimers combined through various lengths of alkyl and poly(ethylene glycol) (PEG) spacer groups as the simple models of multivalent-ligand molecule of GADP and evaluated their immunological enhancement activities in vitro. The GADP dimers showed a higher level stimulatory activities against macrophage-like cells than free GADP and monomeric GADP derivatives. © 1997 Elsevier Science Ltd.