Switching of metabolic-rate scaling between allometry and isometry in colonial ascidians

The metabolic rate and its scaling relationship to colony size were studied in the colonial ascidian Botrylloides simodensis. The colonial metabolic rate, measured by the oxygen consumption rate (V(O2) in millilitres of O(2) per hour) and the colony mass (wet weight M(w) in grams) showed the allometric relationship (V(O2) = 0.0412 M(w)(0.799). The power coefficient was statistically not different from 0.75, the value for unitary organisms. The size of the zooids and the tunic volume fraction in a colony were kept constant irrespective of the colonial size. These results, together with the two-dimensional colonial shape, excluded shape factors and colonial composition as possible causes of allometry. Botryllid ascidians show a takeover state in which all the zooids of the parent generation in a colony degenerate and zooids of a new generation develop in unison. The media for connection between zooids such as a common drainage system and connecting vessels to the common vascular system experienced reconstruction. The metabolic rate during the takeover state was halved and was directly proportional to the colonial mass. The scaling thus changed from being allometric to isometric. The alteration in the scaling that was associated with the loss of the connection between the zooids strongly support the hypothesis that the allometry was derived from mutual interaction among the zooids. The applicability of this hypothesis to unitary organisms is discussed.     

Phenotypic plasticity and morphological integration in a marine modular invertebrate

Background  

Colonial invertebrates such as corals exhibit nested levels of modularity, imposing a challenge to the depiction of their morphological evolution. Comparisons among diverse Caribbean gorgonian corals suggest decoupling of evolution at the polyp vs. branch/internode levels. Thus, evolutionary change in polyp form or size (the colonial module sensu stricto) does not imply a change in colony form (constructed of modular branches and other emergent features). This study examined the patterns of morphological integration at the intraspecific level. Pseudopterogorgia bipinnata (Verrill) (Octocorallia: Gorgoniidae) is a Caribbean shallow water gorgonian that can colonize most reef habitats (shallow/exposed vs. deep/protected; 1–45 m) and shows great morphological variation.     

Branching and self-organization in marine modular colonial organisms: a model

Despite the universality of branching patterns in marine modular colonial organisms, there is neither a clear explanation about the growth of their branching forms nor an understanding of how these organisms conserve their shape during development. This study develops a model of branching and colony growth using parameters and variables related to actual modular structures (e.g., branches) in Caribbean gorgonian corals (Cnidaria). Gorgonians exhibiting treelike networks branch subapically, creating hierarchical mother-daughter relationships among branches. We modeled both the intrinsic subapical branching along with an ecological-physiological limit to growth or maximum number of mother branches (k). Shape is preserved by maintaining a constant ratio (c) between the total number of branches and the mother branches. The size frequency distribution of mother branches follows a scaling power law suggesting self-organized criticality. Differences in branching among species with the same k values are determined by r (branching rate) and c. Species with rr/2 or c>r>0). Ecological/physiological constraints limit growth without altering colony form or the interaction between r and c. The model described the branching dynamics giving the form to colonies and how colony growth declines over time without altering the branching pattern. This model provides a theoretical basis to study branching as a simple function of the number of branches independently of ordering- and bifurcation-based schemes.     

Seasonal patterns of population structure in a colonial marine invertebrate (Bugula stolonifera, Bryozoa)

For sessile invertebrates, the degree to which dispersal mechanisms transport individuals away from their natal grounds can have significant ecological implications. Even though the larvae of the marine bryozoan Bugula stolonifera have limited dispersal potential, high levels of genetic mixing have been found within their conspecific aggregations. In this study, we investigated whether this high mixing within aggregations of B. stolonifera also resulted in high mixing between aggregations. Adult colonies were collected from five sites within and one site outside of Eel Pond, Woods Hole, Massachusetts, in August 2009 and genotyped at 10 microsatellite loci. Significant genotypic differentiation was found between most sites, suggesting limited connectivity across sites, even those separated by only 100 m. This investigation was extended to determine if low levels of genetic mixing throughout the reproductive season could result in increased homogeneity between sites. Four of the five sites in Eel Pond were sampled early, mid-, and late in the reproductive season in 2010, and again in early 2011. Inter- and intra-annual genotypic differentiation was then assessed within and between sites. Results from these analyses document that low levels of mixing could result in increased homogeneity between some aggregations, but that barriers to genetic exchange prevented mixing between most sites. Further, results from inter-annual comparisons within sites suggest that any potential homogeneity achieved throughout the reproductive season will likely be lost by the beginning of the next reproductive season due to the annual cycle of colony die-back and regrowth experienced by B. stolonifera colonies in Eel Pond.     

Allometry of resource capture in colonial cnidarians and constraints on modular growth

1. Indeterminacy in growth of colonial organisms, such as corals, is commonly attributed to their modular construction which frees the colony from the allometric constraints that limit the size of single modules. However, as a colony grows, there may be a decrease in resource availability to interior modules because of active depletion and/or passive deflection by modules on the exterior. The effects of 'self-shading' on resource capture in modular animals are modelled using a simple allometric growth function.
2. The model assumes that resource capture by a module scales as an exponent ( γ ) of colony size (i.e. number of modules). Data taken from the literature indicate that model values of γ for light and prey capture range from – 0·80 to – 1·16 for branching and encrusting corals. Module-specific rates of resource use (i.e. metabolism) are less affected by colony size. Therefore, as a colony grows, net resource state eventually reaches zero, making further growth unsustainable or determinate.
3. The model also predicts an inverse relationship between module size and colony size such as that observed in Caribbean corals. This negative correlation results from the additive effects of module size and colony size on the degree of self-shading.
4. Resource capture is affected by growth form and flow regime, and the interaction between them can account for some of the morphological variation in corals and other colonial suspension feeders.     

The relationship between growth and allometry

The asymptotic allometric equations are derived by considering the relative growth of different physiological and anatomical quantities. The growth rate is assumed to be proportional to the value of the quantity, where the proportion coefficient depends on time. The allometric exponents are calculated for some organs of man and compared with the experimental values for primates. The exponents are generally time-dependent during the growth. The effect of the choice of the origin of time axis on the asymptotic allometric exponents is studied.     

Evolution and dynamics of branching colonial form in marine modular cnidarians: gorgonian octocorals

Multi-branched arborescent networks are common patterns for many sessile marine modular organisms but no clear understanding of their development is yet available. This paper reviews new findings in the theoretical and comparative biology of branching modular organisms (e.g. Octocorallia Cnidaria) and new hypotheses on the evolution of form are discussed. A particular characteristic of branching Caribbean gorgonian octocorals is a morphologic integration at two levels of colonial organization based on whether the traits are at the module or colony level. This revealed an emergent level of integration and modularity produced by the branching process itself and not entirely by the module replication. In essence, not just a few changes at the module level could generate changes in colony architecture, suggesting uncoupled developmental patterning for the polyp and branch level traits. Therefore, the evolution of colony form in octocorals seems to be related to the changes affecting the process of branching. Branching in these organisms is sub-apical, coming from mother branches, and the highly self-organized form is the product of a dynamic process maintaining a constant ratio between mother and daughter branches. Colony growth preserves shape but is a logistic growth-like event due to branch interference and/or allometry. The qualitative branching patterns in octocorals (e.g. sea feathers, fans, sausages, and candelabra) occurred multiple times when compared with recent molecular phylogenies, suggesting independence of common ancestry to achieve these forms. A number of species with different colony forms, particularly alternate species (e.g. sea candelabrum), shared the same value for an important branching parameter (the ratio of mother to total branches). According to the way gorgonians branch and achieve form, it is hypothesized that the diversity of alternate species sharing the same narrow variance in that critical parameter for growth might be the product of canalization (or a developmental constraint), where uniform change in growth rates and maximum colony size might explain colony differences among species. If the parameter preserving shape in the colonies is fixed but colonies differ in their growth rates and maximum sizes, heterochrony could be responsible for the evolution among some gorgonian corals with alternate branching.     

Temperature-mediated stability of the interaction between spider mites and predatory mites in orchards

The nonlinear behavior of the Holling-Tanner predatory-prey differential equation system, employed by R.M. May to illustrate the apparent robustness of Kolmogorov’s Theorem when applied to such exploitation systems, is re-examined by means of the numerical bifurcation code AUTO 86 with model parameters chosen appropriately for a temperature-dependent mite interaction on fruit trees. The most significant result of this analysis is that there exists a temperature range wherein multiple stable states can occur, in direct violation of May’s interpretation of this system’s satisfaction of Kolmogorov’s Theorem: namely, that linear stability predictions have global consequences. In particular these stable states consist of a focus (spiral point) and a limit cycle separated from each other in the phase plane by an unstable limit cycle, all of which are associated with the single community equilibrium point of the system. The ecological implications of such metastability, hysteresis, and threshold behavior for the occurrence of outbreaks, the persistence of oscillations, the resiliency of the system, and the biological control of mite populations are discussed.     

Relationship between current velocity,depth and the invertebrate community in a stable river system

The relationship between invertebrate densities, current velocity and water depth was studied in the Dan River, northern Israel. Maximum current preferences ranged from 5–120 cm sec^–1, and depth preferences ranged from 5–60 cm. Thirty-five taxa of invertebrates were collected by means of colonization cages. Larval and adult stages of 3 Elmidae (Coleoptera) species were treated separately: Limnius letourneuxi, Grouvellinus caucasicus and Elmis rioloides. Differences in current preference were observed between larval and adult stages of the same species of Elmidae. Taxa were also grouped according to preference for turbulence. Wide ranges of depth and current velocity preferences were observed. Most of the taxa were found at between 80–100 cm sec^–1 and at depths of less than 30 cm. A correlation between species diversity and current velocity was established. Velocities of 60–80 cm sec^–1 contained the greatest overlap of faunal preference. The sensitivity of selected species to stream flow reduction is discussed.IES Laboratory, Department of Zoology, Hebrew University; and the Nature Reserves Authority     

Temperature-mediated relationship between western flower thrips (Thysanoptera: Thripidae) and chrysanthemum

Modeling the effect of temperature on the sustainability of insect-plant interactions requires assessment of both insect and plant performance. We examined the effect of temperature on western flower thrips, Frankliniella occidentalis (Pergande), a generalist herbivore with a high reproductive rate, and chrysanthemum inflorescences, a high quality but relatively fixed, ephemeral resource for thrips population growth. We hypothesized that different thrips versus plant responses to temperature result in significant statistical interaction of temperature with thrips abundance and flower damage attributes over time. Experiments were conducted at five temperatures between 20.7 and 35.3 degrees C, with thrips infestation and time after infestation as main effects. Only minor, uncontrolled variations in relative humidity and light intensity may otherwise have influenced the results. High temperatures lead to an initially rapid increase in density of thrips followed by abrupt declines in abundance. The rate of floral senescence increased with temperature and thrips infestation, as indicated by a reduced fresh biomass and greater leaching of yellow pigments. Multiple regression indicated that indices of plant damage responded more directly to thrips density at low than high temperature, supporting the conclusion that temperature affected the outcome beyond what was predictable simply from differential plant and insect optima. The relative intensity of damage caused by individual thrips decreased with increasing temperature, likely caused by thrips competition and reduced survival, growth, and fecundity on depleted inflorescences. Reduced per capita damage at high temperature may be common in insects exploiting fixed plant resources that exhibit an accelerated rate of deterioration at high temperatures.     

Ecology,physiology, allometry and dimensionality

Many ecologically important empirical laws governing physiological rates can be derived from a small set of assumptions. The body size of an organism, the density of tissue and the caloric content of tissue comprise a fundamental set of dimensional quantities. Dimensional arguments can account for a wide variety of ecophysiological observations such as the difference between the size dependence of respiration in terrestrial and aquatic organisms and the independence of net growth efficiency on body size.     

Gompertz curves,allometry and embryogenesis

A number of new properties of the Gompertz curve are derived. In particular, the curve is shown to be of particular importance in the theory of biological similitude, because of a direct relation to the allometric law. A discussion is included indicating some of the practical limitations of the theory.     

Unusual allometry between respiration rate and body size in Chaoborus species

The respiration rates of all four instars of Chaoborus flavicanswere measured with a flow-through respirometer at an experimentaltemperature of 20°C. The respiration rates (µg O₂larva^-1 h^-1) increased parallel to the larval stages accordingto R = 0.027 x W^0.416 (W = µg dry weight), reaching arespiration rate eight times higher for instar IV than for instarI. The slope of the increase with body weight was as low asin two tropical Chaoborus species and was considerably lowerthan usually found for other aquatic animals. Instar IV larvaecollected in the spring showed a significantly higher respirationrate than those collected in the fall. The respiration rateof the fourth instar approximately doubled with a Q₁₀ of 2.1when the experimental temperature was increased from 10 to 20°C.