Flow and polypide distribution in the cheilostome bryozoan Bugula and their inference in Archimedes

Cilia-generated flow in the absence of ambient current is directed from frontal to reverse sides of branches in Bugula turrita, B. turbinata, B. neritina , and B. stolonifera , whether axes of feeding lophophores are perpendicular to the basal plane of branches or are tilted toward distal ends of branches. Ambient current less than 5 cm per second interacts with cilia-generated flow, but ambient flow of 15 cm per second destroys self-generated colonial flow and severely hampers feeding. Polypides are located in the more distal, younger portions of colonies, in species with and without polypide recycling, whereas zooids in the more proximal, older portions are senesced. Presence of feeding polypides in distal but not in proximal portions of the larger spiralled colonies of B. turrita and B. turbinata results in downward, slightly radially directed flow through the colony. The colonial flow passes directly from one whorl to the next-proximal so that water exits from low around the colony perimeter, and a proximally expanding conical stagnant zone occupies the interior of the colony. A substantial percentage of zooecia in distal whorls of well-preserved Archimedes is filled by sediment and inferred to have been occupied by actively feeding polypides. whereas spar-filled zooecia capped by terminal diaphragms were apparently senesced during the latter part of a colony's existence. The capped zooecia constitute an increasing percentage of the total in more proximal whorls. Generally similar colony form and inferred similarity in distribution of current-generating polypides in spiralled colonies of Bugula and in Archimedes suggest that colony-generated flow in Archimedes was similar to that in Bugula , passing downward and then outward, and only through the distal whorls of the colonies.     

Patterns of reporter gene expression in the phase diagram of Bacillus subtilis colony forms.

Factors governing the morphogenesis of Bacillus subtilis colonies as well as the spatial-temporal pattern of expression of a reporter gene during colony development were examined by systematically varying the initial nutrient levels and agar concentrations (wetness), the relative humidity throughout incubation, and the genotype of the inoculum. A relationship between colony form and reporter gene expression pattern was found, indicating that cells respond to local signals during colony development as well as global conditions. The most complex colony forms were produced by motile strains grown under specific conditions such that cells could swim within the colony but not swarm outward uniformly from the colony periphery. The wetness of the growth environment was found to be a critical factor. Complex colonies consisted of structures produced by growth of finger-like projections that expanded outward a finite distance before giving rise to a successive round of fingers that behaved in a similar fashion. Finger tip expansion occurred when groups of cells penetrated the peripheral boundary. Although surfactin production was found to influence similar colony forms in other B. subtilis strains, the strains used here to study reporter gene expression do not produce it. The temporal expression of a reporter gene during morphogenesis of complex colonies by motile strains such as M18 was investigated. Expression arose first in cells located at the tips of fingers that were no longer expanding. The final expression pattern obtained reflects the developmental history of the colony.     

Basis of ontogenetic and evolutionary transformations of thecate hydroids

The high diversity of spatial organization of shoots in colonies of thecate hydroids (Cnidaria, Hydroidomedusa, Leptomedusae) is determined by their modular organization, which is characterized by the cyclic morphogenesis in the colony. It is attempted to show that evolutionary and ontogenetic changes in the spatial organization of hydroids of this group are based on the allometric growth of modules of colony shoots. An increase in size of a developing module provides prerequisites for earlier initiation of the growing tips of succeeding moduls (heterochrony). In some cases, heterochronies determined transition from cyclic to acyclic morphogenesis. The earlier emergence of new growing tips allowed integration of several primary modules into secondary modules, resulting among other things in changes in relative positions of primary modules (heterotopy). In complex colonies, these changes are traced in the ontogeny of a single colony.     

The significances of bacterial colony patterns

Bacteria do many things as organized populations. We have recently learned much about the molecular basis of intercellular communication among prokaryotes. Colonies display bacterial capacities for multicellular coordination which can be useful in nature where bacteria predominantly grow as films, chains, mats and colonies. E. coli colonies are organized into differentiated non-clonal populations and undergo complex morphogenesis. Multicellularity regulates many aspects of bacterial physiology, including DNA rearrangement systems. In some bacterial species, colony development involves swarming (active migration of cell groups). Swarm colony development displays precise geometrical controls and periodic phenomena. Motile E. coli cells in semi-solid media form organized patterns due to chemotactic autoaggregation. On poor media, B. subtilis forms branched colonies using group motility and longrange chemical signalling. The significances of bacterial colony patterns thus reside in a deeper understanding of prokaryotic biology and evolution and in experimental systems for studying self-organization and morphogenesis.     

Scanning electron microscope study of Pseudomonas putida colonies.

Pseudomonas putida colonies were examined by scanning electron microscope. A variety of cell morphologies, multicellular arrangements, and extracellular materials were observed in the fixed material. Different regions of a single colony showed characteristic organizations of these architectural elements. In some cases, the detailed microstructure of the fixed colony surfaces observed by scanning electron microscopy could be correlated with macroscopic patterns visualized by histochemical staining and surface relief photography of live colonies. Extracellular materials were seen to extend onto the agar surface beyond the boundaries of the cell mass, and the final structures of these materials, after fixation and desiccation, were colony specific. The significance of these features of colony microstructure for formulating hypotheses about the control of colony morphogenesis is discussed.     

The Role of Non-Foraging Nests in Polydomous Wood Ant Colonies

A colony of red wood ants can inhabit more than one spatially separated nest, in a strategy called polydomy. Some nests within these polydomous colonies have no foraging trails to aphid colonies in the canopy. In this study we identify and investigate the possible roles of non-foraging nests in polydomous colonies of the wood ant Formica lugubris. To investigate the role of non-foraging nests we: (i) monitored colonies for three years; (ii) observed the resources being transported between non-foraging nests and the rest of the colony; (iii) measured the amount of extra-nest activity around non-foraging and foraging nests. We used these datasets to investigate the extent to which non-foraging nests within polydomous colonies are acting as: part of the colony expansion process; hunting and scavenging specialists; brood-development specialists; seasonal foragers; or a selfish strategy exploiting the foraging effort of the rest of the colony. We found that, rather than having a specialised role, non-foraging nests are part of the process of colony expansion. Polydomous colonies expand by founding new nests in the area surrounding the existing nests. Nests founded near food begin foraging and become part of the colony; other nests are not founded near food sources and do not initially forage. Some of these non-foraging nests eventually begin foraging; others do not and are abandoned. This is a method of colony growth not available to colonies inhabiting a single nest, and may be an important advantage of the polydomous nesting strategy, allowing the colony to expand into profitable areas.     

Self-similar colony morphogenesis by gram-negative rods as the experimental model of fractal growth by a cell population.

The ability to form a fractal colony was shown to be common among several species of the family Enterobacteriaceae. Bacterial spreading growth in a two-dimensional field of nutrient concentration was indicated to be important for this experimental self-similar morphogenesis. As a basic analogy, the diffusion-limited aggregation model was suggested. Fractal dimensions of colonies were mostly in the range of values from 1.7 to 1.8, similar to those of the two-dimensional diffusion-limited aggregation model. Bacterial characteristics and culture conditions inducing changes in fractal patterns and growth rates were identified. The contribution of the bacterial multicellular nature to fractal morphogenesis is discussed.     

Biological attributes of colony-type variants of Candida albicans

Twenty 'commensal' oral or 'pathogenic' vaginal isolates of Candida albicans were examined for colony morphology on malt/yeast-extract and serum-based agar media. Diverse and variable colony morphology was seen on serum agar. In 17 strains, selective subculture of morphologically atypical colonies produced progeny which had reverted to the morphology of the majority of parental colonies. However, in one strain, a highly stable colony variant was isolated which did not revert on subculture. In two further strains, variants were isolated which could be maintained with at least 99% homogeneous colony type by selective colony subculture, but reversion to the parental type or switching to other morphologies occurred at rates of 10(-2) to 10(-4): a rapid switching phenomenon. The relative proportions of mycelial or yeast forms were the main determinants of colony morphology. The variants were biotyped using a selection of biochemical tests. The stable variant differed from its parent in several characters, including rate of production of a proteinase enzyme. The pathogenicity of variants was compared in mice, and both stable and switching variants differed in virulence from their parental strains. Colony-type variation on suitable media is thus a powerful tool in the isolation of mutants or variants of C. albicans which differ from 'isogenic' parents in significant biological properties. Such variants may aid identification and characterization at the molecular level of determinants of, for example, pathogenicity and morphogenesis.     

Self-similar colony morphogenesis by gram-negative rods as the experimental model of fractal growth by a cell population.

The ability to form a fractal colony was shown to be common among several species of the family Enterobacteriaceae. Bacterial spreading growth in a two-dimensional field of nutrient concentration was indicated to be important for this experimental self-similar morphogenesis. As a basic analogy, the diffusion-limited aggregation model was suggested. Fractal dimensions of colonies were mostly in the range of values from 1.7 to 1.8, similar to those of the two-dimensional diffusion-limited aggregation model. Bacterial characteristics and culture conditions inducing changes in fractal patterns and growth rates were identified. The contribution of the bacterial multicellular nature to fractal morphogenesis is discussed.     

The influence of the vibration signal on worker interactions with the nest and nest mates in established and newly founded colonies of the honey bee, Apis mellifera

Honey bees adjust cooperative activities to colony needs, based in part on information acquired through interactions with the nest and nest mates. We examined the role of the vibration signal in these interactions by investigating the influence of the signal on the movement rates, cell inspection activity, and trophallaxis behavior of workers in established and newly founded colonies of the honey bee, Apis mellifera. Compared to non-vibrated control bees, vibrated recipients in both colony types exhibited increased movement through the nest and greater cell inspection activity, which potentially increased contact with stimuli that enhanced task performance. Also, compared to controls, recipients in both colony types showed increased rates of trophallactic interactions and spent more time engaged in trophallaxis, which potentially further increased the acquisition of information about colony needs. The vibration signal may therefore help to organize labor in honey bees in part by increasing the rate at which workers obtain information about their colony. Vibrated recipients in the established and newly founded colonies did not differ in any aspect of behavior examined, suggesting that colony developmental state did not influence the degree to which individual workers responded to the signal. However, previous work has demonstrated that newly founded colonies have increased levels of vibration signal behavior. Thus, the vibration signal may help to adjust worker activity to colony conditions partly by stimulating greater numbers of bees to acquire information about colony needs, rather than by altering the level at which individual recipients react to the signal. Received 23 October 2006; revised 15 January 2007; accepted 7 February 2007.     

Sex ratios and the distribution of elaiosomes in colonies of the ant, Aphaenogaster rudis

Summary: Genetic theory predicts that workers in monogynous ant colonies with singly-mated queens should capitalize on higher relatedness with sisters than with brothers by altering the sex investment ratio of a colony in favor of females. Sex investment ratios, however, may also be influenced by the amount of resources available to colonies, in part because more mating opportunities might be obtained by investing scarce resources in males, which are much smaller than queens. Female larvae that reach a critical size by a particular point in development become queens while underfed larvae develop into workers, so workers could potentially influence the sex investment ratio of a colony by selectively feeding female larvae. In a previous experiment on the ant, Aphaenogaster rudis, colonies increased female sex investment after their diet was supplemented with elaiosomes, a lipid-rich food gained from a seed dispersal mutualism. In order to investigate the mechanisms producing this shift, we radio-labeled Sanguinaria canadensis elaiosomes with fatty acids and compared uptake among castes within a colony. The experiment was performed in both the laboratory and field. Lab colonies produced female-biased sex investment ratios, while field colonies mainly invested in males. We hypothesize that this discrepancy is related to differing levels of background food availability in the lab and field. The results of the elaiosome distribution experiment do not support a hypothesis that elaiosomes play a qualitative role in queen determination, because all individuals in a colony receive this nutrient. There is, however, support for the hypothesis that elaiosomes have a quantitative effect on larval development because larvae that accumulated more radio-label from elaiosomes tended to develop into gynes (virgin queens), while other female larvae developed into workers.     

COLONY FORMATION AND SEXUAL MORPHOGENESIS IN THE COCCOLITHOPHORE PLEUROCHRYSIS SP. (HAPTOPHYTA)1

Pleurochrysis sp. formed two types of symmetrical, diploid colonies on solid media: (i) single‐cell lineage (SCL) colonies and (ii) aggregation (AG) colonies. The first division of a single mother cell was asymmetric in ～54% of SCL colonies. These colonies developed at a slower rate than AG colonies. Diffusible molecules released from the cells acted like morphogens enhancing formation of AG colonies; their influence on chemotaxis of aggregating cells was dependent on concentration of the inoculum. Nitrogen depletion of diploid colonies induced sexual morphogenesis and colony patterning into inner and outer regions. The smaller innermost cells were surrounded by outer larger cells. Developmental mechanisms of colony formation were examined in relation to the heteromorphic, haplo‐diploid life cycle.     

Asymptotic analysis of a chemotactic model of bacteria colonies

An estimate of the distance between spots generated by a bacterial colony model is obtained. The model describes the morphogenesis of a spot pattern in colonies of chemotactic strains of Escherichia coli. Asymptotic methods for other cell-chemotaxis models, which have been successfully used by previous researchers, can be applied also to this model. However the calculations and the result is more complicated for this model. The result is verified by comparing it with the results by numerical computations of solutions of the model.     

The economics of brood raiding and nest consolidation during ant colony founding

Summary The most dangerous time for an ant colony is during the founding stage when the small colony is vulnerable to predation and competition. Colonies can grow more rapidly when multiple queens cooperate in raising the first worker brood (pleometrosis) or by raiding other incipient colonies for their brood. This brood raiding has been proposed to be the primary force selecting for pleometrosis, i.e. multiple-queen colonies may have a considerable advantage in destroying neighbours by aggressively stealing their brood. An alternative hypothesis is that incipient nests are part of a larger, interconnected population structure and that brood raiding reflects cooperative pleometrosis with subdivided colonies. A simple mathematical model supports the second hypothesis: workers of incipient colonies are especially favoured to peaceably abandon their nest and join with other colonies if the queens are related or queens from raided colonies can infiltrate the raiding colony. The latter condition is often met in ant species that brood raid and particularly exemplified in fire ants (Solenopsis invicta), where brood raiding involves little mortal combat and combines with pleometrosis to rapidly increase colony size. It is proposed that the term nest consolidation should replace brood raiding to more accurately reflect the relatively non-aggressive and potentially apparently cooperative nature of interactions between incipient ant colonies.     

Spatial differentiation in the vegetative mycelium of Aspergillus niger

Fungal mycelia are exposed to heterogenic substrates. The substrate in the central part of the colony has been (partly) degraded, whereas it is still unexplored at the periphery of the mycelium. We here assessed whether substrate heterogeneity is a main determinant of spatial gene expression in colonies of Aspergillus niger. This question was addressed by analyzing whole-genome gene expression in five concentric zones of 7-day-old maltose- and xylose-grown colonies. Expression profiles at the periphery and the center were clearly different. More than 25% of the active genes showed twofold differences in expression between the inner and outermost zones of the colony. Moreover, 9% of the genes were expressed in only one of the five concentric zones, showing that a considerable part of the genome is active in a restricted part of the colony only. Statistical analysis of expression profiles of colonies that had either been or not been transferred to fresh xylose-containing medium showed that differential expression in a colony is due to the heterogeneity of the medium (e.g., genes involved in secretion, genes encoding proteases, and genes involved in xylose metabolism) as well as to medium-independent mechanisms (e.g., genes involved in nitrate metabolism and genes involved in cell wall synthesis and modification). Thus, we conclude that the mycelia of 7-day-old colonies of A. niger are highly differentiated. This conclusion is also indicated by the fact that distinct zones of the colony grow and secrete proteins, even after transfer to fresh medium.     

Morphology of Ureaplasma urealyticum (T-mycoplasma) organisms and colonies.

The morphology of Ureaplasm urealyticum in broth cultures was studied by phase-contrast microscopy. Most organisms appeared singly or in pairs. Long filaments and long chains of cocci, common in classical mycoplasma cultures, were not observed. On solid medium, U. urealyticum produced "fried-egg" colonies which developed according to the scheme suggested by Razin and Oliver (J. Gen. Microbiol., 1961) for the morphogenesis of the classical mycoplasma colonies. The formation of the peripheral zone of the colonies followed that of the central zone only when growth conditions were adequate, Hence, the appearance of peripheral zones, and consequently the larger colony size, can be taken as an indicator of improved growth conditions. Incubation in an atmosphere of 100% CO2 resulted in significantly larger colonies than in an atmosphere of N2, O2, or air. CO2 acts as a buffer, keeping the pH at the optimal range for Ureaplasma growth (pH 6.0 to 6.5) in the presence of the ammonia produced from the urea hydrolyzed by the organisms. The addition to the medium of 0.01 M urea together with 0.01 M putrescine enabled better growth than with urea alone. Small amounts of phosphate improved growth in an atmosphere of CO2, apparently fulfilling a nutritional role. Under nitrogen, higher phosphate concentrations were required for good growth, apparently serving as a buffer as well as a nutrient. Sodium chloride and sucrose which had been added to increase the tonicity of the medium inhibited growth above 0.1 M. An increase in the agar concentration above 2% resulted in decreased colony size. Likewise, prolonged drying of the agar plates caused a marked decrease in colony size, mostly affecting the peripheral zone. The addition of both urea and putrescine to the growth medium and incubation in a humidified CO2 atmosphere are recommended for improved growth and formation of fried-egg colonies of U. ureaplyticum on agar. It must be emphasized that these experiments were carried out with a laboratory-adapted strain.     

The spatial distribution and size of rook (Corvus frugilegus) breeding colonies is affected by both the distribution of foraging habitat and by intercolony competition

Explanations for the variation in the number of nests at bird colonies have focused on competitive or habitat effects without considering potential interactions between the two. For the rook, a colonial corvid which breeds seasonally but forages around the colony throughout the year, both the amount of foraging habitat and its interaction with the number of competitors from surrounding colonies are important predictors of colony size. The distance over which these effects are strongest indicates that, for rooks, colony size may be limited outside of the breeding season when colony foraging ranges are larger and overlap to a greater extent.     

A stochastic model of self-renewal and commitment to differentiation of the primitive hemopoietic stem cells in culture

We recently identified a murine hemopoietic stem cell colony which consists of undifferentiated (blast) cells and appears to be more primitive than CFU-GEMM in the stem cell hierarchy. The progenitors for the colony which we termed “stem cell colony” possess an extensive self-renewal capacity and the ability to generate many secondary multipotential hemopoietic colonies in culture. We replated a total of 68 stem cell colonies from cultures of murine spleen cells and analyzed the number of stem cell–and granulocyte(neutrophil)-erythrocyte-macrophage-megakaryocyte (GEMM) colony-forming cells in individual stem cell colonies. Of the 68 stem cell colonies, 35 contained progenitors (abbreviated as “S”-cells) for stem cell colonies. The distributions of S-cells and CFU-GEMM in individual stem cell colonies were extremely heterogeneous. Neither the frequency distributions of S-cells nor CFU-GEMM in stem cell colonies could be fitted well by Poisson distribution. Rather, the frequency distribution of the s-cells could be approximated by a geometric distribution and that of CFU-GEMM by an exponential distribution, both of which are variates of the gamma distribution. Our observations are in agreement with those on the distributions of CFU-S in individual spleen colonies and provided support for a stochastic model for stem cell self-renewal and commitment in culture. Application of the theory of the branching process to the distribution of S-cells revealed a distributional parameter “p” of 0.589 which is also in agreement with the earlier report on the p value for reproduction of CFU-S.