Observations of the tissue-skeleton interface in the scleractinian coral <Emphasis Type="Italic">Stylophora pistillata</Emphasis>

Recent micro-analytical studies of coral skeletons have led to the discovery that the effects of biology on the skeletal chemical and isotopic composition are not uniform over the skeleton. The aim of the present work was to provide histological observations of the coral tissue at the interface with the skeleton, using Stylophora pistillata as a model, and to discuss these observations in the context of skeletal ultra-structural organization and composition. Several important observations are reported: (1) At all scales of observation, there was a precise morphological correspondence between the tissues and the skeleton. The morphological features of the calicoblastic ectoderm correspond exactly to the shape of individual crystal fiber bundles in the underlying skeleton, indicating that the calicoblastic cell layer is in direct physical contact with the skeletal surface. This is consistent with the previously observed chemical and isotopic composition of the ultra-structural components in the skeleton. (2) The distribution and density of desmocyte cells, which anchor the calicoblastic ectoderm to the skeletal surface, vary spatially and temporally during skeletal growth. (3) The tissue above the coenosteal spines lack endoderm and consists only of ectodermal cell-layers separated by mesoglea. These findings have important implications for models of vital effects in coral skeletal chemistry and isotope composition.     

Diurnal patterns of skeleton formation in Pocillopora damicornis (Linnaeus)

Scanning, transmission and X-ray microanalytical electron microscopy were used to investigate the skeleton, organic matrix and calicoblastic ectoderm of the reef coral Pocillopora damicornis over a diurnal cycle. All skeletal surfaces, both during day and hight, are fasciculate except for skeletal spines on the branch tip apex at night where small (0.5 m) fusiform crystals are deposited. X-ray microanalysis shows that the fusiform crystals and needle-shaped crystals that compose the fasciculi are distinct forms of calcium carbonate. Demineralization of the skeleton reveals an organic matrix with two components which are related to the formation of fusiform crystals and fasciculi. During the day the calicoblastic ectoderm overlying all skeletal surfaces is 1–3 m thick. At night ultrastructural evidence suggests that skeletal deposition occurs only on those skeletal spines at the branch tip apex which are growing parallel with the branch growth axis. The calicoblastic ectoderm overlying apical skeletal spines at night shows a greater degree of cellular activity, and is thicker, than calicoblastic ectoderm overlying both other skeletal surfaces at night (<8 m cf. >6 m) and all skeletal surfaces during the day (<8 m cf. >3 m). The deposition of fusiform crystals on skeletal spines at the branch tip apex is proposed to promote deposition of fasciculi during the day, relative to other skeletal surfaces, providing a mechanism determining apical growth of branch tips. The results are discussed with respect to previous concepts of skeletal deposition in scleractinian corals.     

Low temperature FESEM of the calcifying interface of a scleractinian coral

The ultrastructural nature of the calcifying interface in the scleractinian coral Galaxea fascicularis has been investigated using high-resolution, low temperature field emission scanning electron microscopy (FESEM). This technique permitted structural analyses of soft tissue and skeleton in G. fascicularis in a frozen-hydrated state, without the need for chemical fixation or decalcification. Structural comparisons are made between frozen-hydrated polyps and polyps that have undergone conventional fixation and decalcification. Vesicles expelled by the calicoblastic ectodermal cells into sub-skeletal spaces and previously suggested to play a role in calcification were commonly observed in fixed samples but were distinctly absent in frozen-hydrated preparations. We propose that these vesicles are fixation artefacts. Two distinct types of vesicles (380 and 70 nm in diameter, respectively), were predominant throughout the calicoblastic ectodermal cells of frozen-hydrated preparations, but these were never seen to be entering, or to be contained within, sub-skeletal spaces, nor did they contain any crystalline material. In frozen-hydrated preparations, membranous sheets were seen to surround and isolate portions of aboral mesogloea and to form junctional complexes with calicoblastic cells. The calicoblastic ectoderm was closely associated with the underlying skeleton, with sub-skeletal spaces significantly smaller (P<0.0001) in frozen-hydrated polyps compared to fixed polyps. A network of organic filaments (26 nm in diameter) extended from the apical membranes of calicoblastic cells into these small sub-skeletal cavities. A thin sheath was also frequently observed adjacent to the apical membrane of calicoblastic cells.     

Calcium localisation by X-ray microanalysis and fluorescence microscopy in larvae of zooxanthellate and azooxanthellate corals

X-ray microanalysis and fluorescence microscopy (Calcium Orangetrade mark) was used to determine the distribution of intracellular calcium (I(Ca)), in the form of total and ionic calcium respectively, in planulae and settled larvae of a zooxanthellate coral. The distribution of total calcium only was determined in larvae of an azooxanthellate coral. In azooxanthellate planulae and settled larvae, total I(Ca) concentration in the oral ectoderm was high and similar to that in seawater (SW). Calcium concentration did not vary (P > 0.05) between planulae and settled larvae. However, settled larvae accumulated large amounts of calcium in gastrodermal lipid-containing cells. In contrast, zooxanthellate planulae possessed significantly (P < 0.01) lower concentrations of total I(Ca) within ectodermal cells in comparison to settled larvae. In addition, in settled zooxanthellate larvae total calcium concentration in the mesogloea and coelenteron was significantly (P < 0.05) higher than in the oral ectodermal and gastrodermal cells, respectively. Total I(Ca) concentrations in the oral ectoderm of settled larvae were also significantly (P < 0.01) lower than that of the calicoblastic ectoderm. In zooxanthellate settled larvae, ionic I(Ca) levels in the aboral epithelium surrounding rapidly growing septa were high. These levels increased significantly (P < 0.05) within the tissue surrounding growing septa after incubation in high-calcium SW.     

The role of polyp-stolon junctions in the redox signaling of colonial hydroids

An encrusting colonial hydroid can be regarded as a network of polyps or ‘mouths’ connected by tube-like stolons. The success of the colony crucially depends on putting these mouths where the available food is. Feeding-related perturbations may provide important signals in this regard. After feeding, polyps contract regularly, dispersing food throughout the colony via the gastrovascular fluid. Mitochondrion-rich epitheliomuscular cells concentrated near polyp-stolon junctions likely drive these contractions. Putatively, the redox state of these cells may influence colony-level form. For instance, the metabolic demand associated with feeding-related contractions results in mitochondria that have relatively oxidized electron carriers and produce lesser amounts of reactive oxygen species (ROS). ROS or other redox-sensitive molecules emitted from polyp-stolon junctions into the gastrovascular fluid may provide stolons with signals influencing elongation, branching, and regression. Treatments of colonies with anti-oxidants cause peripheral stolon tips to rapidly regress. This regression appears to be an active process involving a flux of locally produced peroxides and cell and tissue death. At the same time, polyps and stolon tips in the center of treated colonies remain healthy. ‘Sheet-like’ growth of short, branched stolons ensues. Signals that inhibit the outward growth of stolons may lead by default to the concentrated growth of stolons and polyps in food-rich areas. ROS may mediate signaling mechanisms involving nitric oxide, programmed cell death, a variety of redox-regulated proteins, or all of these.     

Propagation of the threatened staghorn coral Acropora cervicornis: methods to minimize the impacts of fragment collection and maximize production

Coral reef restoration methods such as coral gardening are becoming increasingly considered as viable options to mitigate reef degradation and enhance recovery of depleted coral populations. In this study, we describe several aspects of the coral gardening approach that demonstrate this methodology is an effective way of propagating the threatened Caribbean staghorn coral Acropora cervicornis: (1) the growth of colonies within the nursery exceeded the growth rates of wild staghorn colonies in the same region; (2) the collection of branch tips did not result in any further mortality to the donor colonies beyond the coral removed for transplantation; (3) decreases in linear extension of the donor branches were only temporary and donor branches grew faster than control branches after an initial recovery period of approximately 3–6 weeks; (4) fragmentation did not affect the growth rates of non-donor branches within the same colony; (5) small branch tips experienced initial mortality due to handling and transportation but surviving tips grew well over time; and (6) when the growth of the branch tips is added to the regrowth of the fragmented donor branches, the new coral produced was 1.4–1.8 times more than new growth in undisturbed colonies. Based on these results, the collection of small (2.5–3.5 cm) branch tips was an effective propagation method for this branching coral species resulting in increased biomass accumulation and limited damage to parental stocks.     

Studies on hydroid differentiation. VII. The hydrozoan stolon

The stolon of the colonial marine hydroid Podocoryne carnea differentiates sequentially as a function of age, forming four distinguishable regions characterized by epidermal cell differentiation: The Tip, New Stolon, Cnidogenic Masses, Old Stolon. Radioautographs of sections of colonies exposed to tritiated thymidine show that although cells of the epidermis and gastrodermis of the stolon incorporate the nucleoside into acid stable polynucleotide, cells of the stolon tips do not. Stolon extension is not, therefore, the result of a localized meristem-like growth zone. Stolon branching and new polyp formation are, similarly, not signaled by increased thymidine incorporation. The initial event heralding these morphogenetic activities appears to be the reorientation of epidermal cells along a new axis, and the acquisition of perisarc dissolving ability. This evidence is contraindicative of direct dependence of colony form on colony growth. The larger part of stolon epidermal cells are organized into cnidogenic masses where cnidocytes and possibly other amoebocytic cells are produced. Although no mitotic figures have been observed in gastroderm cells of the stolon, thymidine incorporation in this tissue occurs with the same frequency as it does in epidermis. Considerable numbers of gastroderm cells can be found in the gastric cavity. Frequently these and gastroderm cells in the stolon and polyps contain more than one nucleus.     

Holocene coral reef accretion in Hawaii: a function of wave exposure and sea level history

 In the high Hawaiian Islands, significant accretion due to coral reef growth is limited by wave exposure and sea level. Holocene coral growth and reef accretion was measured at four stations off Oahu, Hawaii, chosen along a gradient in wave energy from minimum to maximum exposures. The results show that coral growth of living colonies (linear extension) at optimal depths is comparable at all stations (7.7–10.1 mm/y), but significant reef accretion occurs only at wave sheltered stations. At wave sheltered stations in Hanauma Bay and Kaneohe Bay, rates of long term reef accretion are about 2.0 mm/y. At wave exposed stations, off Mamala Bay and Sunset Beach, reef accretion rates are virtually zero in both shallow (1 m) and deeper (optimal) depths (12 m). At wave sheltered stations, such as Kaneohe Bay and Hanauma Bay, Holocene reef accretion is on the order of 10–15 m thick. At wave exposed stations, Holocene accretion is represented by only a thin veneer of living corals resting on antecedent Pleistocene limestone foundations. Modern coral communities in wave exposed environments undergo constant turnover associated with mortality and recruitment or re-growth of fragmented colonies and are rarely thicker than a single living colony. Breakage, scour, and abrasion of living corals during high wave events appears to be the major source of mortality and ultimately limits accretion to wave sheltered environments. Depth is particularly important as a modulator of wave energy. The lack of coral reef accretion along shallow open ocean coastlines may explain the absence of mature barrier reefs in the high Hawaiian Islands. Accepted: 14 May 1998     

Ontogenetic changes in the capacity of the coral Pocillopora damicornis to originate branches

Most colonial corals vary intraspecifically in growth forms, and the diversity in branching morphology is especially striking. While the effects of environmental factors on growth forms have been studied, the genetic control of coral branching patterns has received little attention. The discovery of ontogenetic changes in the capacity to originate branching would set the stage for studies of how branch formation is genetically controlled. During experiments investigating contact reactions in the coral Pocillopora damicornis, we observed that young colonies derived from settled planulae and colonies regenerated from adult branch tips assumed different growth forms. Young colonies formed at least one branch from the central region of the colony, while colonies regenerated from adult branch tips (3-5 mm long) did not form branches during the 9-month observation period. This pattern was invariable, regardless of the types and outcomes of the contact experiments or the orientation of the branch tips. However, some fragments taken from 1- or 2-year-old colonies formed branches. This suggests that the rate of branch formation in P. damicornis colonies decreases with age. These findings will facilitate investigations of the mechanism of coral branch formation at the molecular level.     

SCENEDESMUS MORPHOGENESIS. TRACE ELEMENTS AND SPINE FORMATION1

When Scenedesmus culture 16, a soil organism with numerous long spines, was grown in axenic culture in a dilute laboratory medium (with approximately 30 mg/liter total salts), spines were not formed. In this medium the organism resembled S. bijugatus, whereas S. longus and S. abundans colonies were typically formed in more concentrated media. Spines were formed on daughter colonies upon transfer to the dilute medium plus additional Fe EDTA or Ca EDTA. No other individual components of the medium did this. Spineless colonies were healthy and green with a well-defined chloroplast, provided the culture was transferred often. In addition to the absence of spines, cultures in the dilute medium had a linear arrangement of cells within a colony, along with terminal cells 20% shorter than median cells. With 4-celled spiny colonies all cells were the same length and were alternately arranged.     

Influence of water flow on skeletal isotopic composition in the coral Pocillopora damicornis

Fragments of branching Pocillopora damicornis coral colonies were grown in experimental flumes under two water flow regimes. Colony size and buoyant weight increased most rapidly in the fast-flow regime. Branch tips from the upper and outer parts of the colonies showed the lowest and most consistent skeletal oxygen isotope ratios. Flow regime had little influence on the lowest oxygen isotope ratios, which were at least 3.5‰ lighter than the calculated oxygen isotopic equilibrium. These “kinetic” isotope effects are comparable to those observed in Porites corals. Relatively more branch tips showed extreme ¹⁸O depletions under low-flow conditions, and among small coral colonies. Isotopic variability was greater among branch tips from the lower and inner parts of the colonies and at high flow. Skeletal oxygen and carbon isotope ratios generally showed positive correlations. Despite the particularly large offsets from isotopic equilibrium, the isotopically lightest branches showed the greatest isotopic consistency and therefore would make the best isotopic thermometers. Isotopic variability within the colony may provide an indication of flow regime.     

THE ROLE OF UNICELLS IN THE POLYMORPHIC SCENEDESMUS ARMATUS (CHLOROPHYCEAE)1

The UTEX 2193 strain of Scenedesmus armatus (Chod.) Chod, when cultured in any of several media (whether natural or artificial, concentrated or dilute) produced a variety of colonial morphologies as well as a unicell population. Morphological expression was related to culture ape. When the initial cell density was just a feu1 hundred cells per mL. the culture first produced a unicell population, then spiny colonies, and as stationary phase was approached, spine-less colonies. Two classes of spiny colonies were detected. Type I colonies had elongate cells with the terminal cells shorter than median cells. Spines were longer than cell length. The wider, oval, grainy cells of Type II colonies were uniform m length. Spines were shorter and thicker than those on Type I colonies. Only Type I colonies produced unicells: the latter appeared as two morphs. The smaller unicell was obovoid with four delicate spines: the larger had ovate cells bearing four thicker spines. Control of unicell development in all media was achieved by carefully monitoring colony type and cell number used for the inoculum. A unicellular population developed in batch culture in defined media, both concentrated and dilute, when the initial cell density (either Type I or Type II colonies) was low (below 1000 cells-mL^?1), as well as in synchronous cultures. With higher initial cell densities, e.g. 2 × 10⁴ cells·mL^?1, the inoculum had to contain Type I colonies to produce unicells. Unicells were also produced in water from Agronomy Pond, where the strain originated. We discuss the role of unicell populations in the distribution of Scenedesmus.     

Structure and function of clypeasteroid miliary spines (Echinodermata,Echinoides)

Summary Histological and ultrastructural techniques have been used to describe the functional morphology of clypeasteroid miliary spines, with special reference to their supposed mucus-secreting role. Mucus cells were not found in the miliary spines of any members of the Arachnoididae, Fibulariidae, Laganidae, Echinarachniidae, Dendrasteridae, Astriclypeidae, or Mellitidae examined in this study. Only members of the Clypeasteridae have mucus-secreting cells in these spines. Characteristics of the skeleton, ultrastructure of the nervous system, and histology of the musculature and epithelia of the base, shaft and tip are also discussed. Miliary spines have two bands of cilia running along the entire length of opposite sides of the shaft. The geometric packing of cilium-bearing cells in these bands is described for the first time, as is the remarkable form of the sacs found at the tips of dendrasterid, astriclypeid, and mellitid miliary spines. These sacs are definitely not mucous sacs, as previously described, but are balloons of single-celled epithelium internally tethered to the skeletal tip by copious quantities of collagenous connective tissue. Miliary spines prevent obstruction of aboral nutritive and ventilatory ciliary currents caused by substrate particles falling to the test surface during burrowing. They do this in two ways: (1) they help generate ciliary currents that sweep finer material off the test, and (2) they contribute to the formation of a spine canopy that mechanically blocks larger particles from falling between the spines. Members of the Clypeasteridae secrete an interspine mucous tent that traps potentially clogging material. The miliary spine sacs of sand dollars are deformable space-fillers that plug holes between primary spines in the aboral canopy, even as the spines rock on their tubercles to push sand backwards over the test. Allometry of spines from Echinarachnius parma suggests that aboral military spines and club-shaped spines exhibit co-ordinated growth that maintains the aboral canopy throughout post-metamorphic ontogeny, and that aboral spins have an overall lower growth rate than spines on the oral surface.     

Cell proliferation and growth inZoothamnium niveum (Oligohymenophora, Peritrichida) — Thiotrophic bacteria symbiosis

Chemolithoautotrophic, sulphide-oxidizing (thiotrophic) symbioses represent spectacular adaptations to fluctuating environmental gradients and survival is often accomplished when growth is fuelled by sufficient nourishment through the symbionts leading to fast cell proliferation. Here we show 5′-bromo-2′ deoxyuridine (BrdU) pulse labelling of vegetative growingZoothamnium niveum, a colonial ciliate obligately associated with thiotrophic ectosymbionts, and demonstrate age related growth profiles in three heteromorphic host cell types. At the colony’s apex, a large top terminal zooid performed high proliferation activity, which decreased significantly with increasing colony age but was still present in old colonies indicating that this cell possesses lifelong cell division potential. In contrast, terminal branch zooids proliferated independent of colony age but appeared to be limited by their cell division capacity predetermined by branch size, thus leading to the strict, feather-shaped colony form. Appearance of labelled terminal branch zooids allowed us to distinguish a highly proliferating apical colony region from an almost inactive, senescent basal region. In macrozooids attached to the colony, extensive BrdU labelling suggests that DNA synthesis occurs in preparation for a new generation. As motile swarmers, the macrozooids seem to be arrested in the cell cycle and mitosis and cell division occur when the swarmer settles and transforms into a top terminal zooid buildingup a new colony.     

Critical Evaluation of Branch Polarity and Apical Dominance as Dictators of Colony Astogeny in a Branching Coral

The high morphological resemblance between branching corals and trees, can lead to comparative studies on pattern formation traits, best exemplified in plants and in some cnidarians. Here, 81 branches of similar size of the hermatypic coral Stylophora pistillata were lopped of three different genets, their skeletons marked with alizarin red-S, and divided haphazardly into three morphometric treatment groups: (I) upright position; (II) horizontal position, intact tip; and (III) horizontal position, cut tip. After 1 y of in-situ growth, the 45 surviving ramets were brought to the laboratory, their tissues removed and their architectures analyzed by 22 morphological parameters (MPs). We found that within 1 y, isolated branches developed into small coral colonies by growing new branches from all branch termini, in all directions. No architectural dissimilarity was assigned among the three studied genets of treatment I colonies. However, a major architectural disparity between treatment I colonies and colonies of treatments II and III was documented as the development of mirror structures from both sides of treatments II and III settings as compared to tip-borne architectures in treatment I colonies. We did not observe apical dominance since fragments grew equally from all branch sides without documented dominant polarity along branch axis. In treatment II colonies, no MP for new branches originating either from tips or from branch bases differed significantly. In treatment III colonies, growth from the cut tip areas was significantly lower compared to the base, again, suggesting lack of apical dominance in this species. Changes in branch polarity revealed genet associated plasticity, which in one of the studied genets, led to enhanced growth. Different genets exhibited canalization flexibility of growth patterns towards either lateral growth, or branch axis extension (skeletal weight and not porosity was measured). This study revealed that colony astogeny in S. pistillata is a regulated process expressed through programmed events and not directly related to simple energy trade-off principles or to environmental conditions, and that branch polarity and apical dominance do not dictate colony astogeny. Therefore, plasticity and astogenic disparities encompass a diversity of genetic (fixed and flexible) induced responses.     

Branching patterns, generating rules, and astogenetic trajectories in arborescent bryozoans

The terminal growth of bryozoans allows a reconstruction of their growth history and generating rules from the developed pattern. Three species of arborescent bryozoans share a bias in growth rate that favors reverser branches (those whose direction of growth is opposite that of their parent branch); this bias produces a common hummocky appearance to the top margin of the colony. Other differences in the growth rules, however, result in markedly different colony forms. For two species, the cue for splitting of branches seems to be the length of the branch; for the other, the cue seems to be the time since the last splitting. Simulations based on the distinction between reversing and nonreversing branches reproduce the hummocky pattern, but offer little insight into the underlying mechanism. Simulations based on occlusion, the line-of-sight blocking of the branches by each other, give similar patterns with fewer assumptions. These simulations also suggest that the pattern is due to a limited availability of nutrients to occluded growing tips, and that nutrients remain relatively localized within colonies. The actual determinants of colony form are likely to combine both nutrition and geometry.     

Aggressive and Docile Colony Defence Patterns in Apis mellifera. A Retreater–Releaser Concept

Colony defence in Apis mellifera involves a variety of traits ranging from ‘aggressive’ (e.g. entrance guarding, recruitment of flying guards) to ‘docile’ (e.g. retreating into the nest) expression. We tested 11 colonies of three subspecies (capensis, scutellata, carnica) regarding their defensiveness. Each colony was selected as reportedly ‘aggressive’, ‘intermediate’ or ‘docile’ and consisted of about 10,000 bees. We applied three stimulation regimes (mechanical disturbance, exposure to alarm pheromones, and the combination of both) and measured their behaviours by tracking the rates of outflying bees at the entrance sites of the test hives. We provided evidence that for mechanical disturbances the test colonies resolved into two response types, if the ‘immediate’ defence response, assessed in the first minute of stimulation, was taken as a function of foraging: ‘releaser’ colonies allocated flying guards, ‘retreater’ colonies reduced the outside-hive activities. This division was observed irrespective of the subspecies membership and maintained in even roughly changing environmental conditions. However, if pheromone and mechanical stimulation were combined, the variety of colony defensiveness restricted to two further types irrespective of the subspecies membership: six of nine colonies degraded their rate of flying defenders with increasing foraging level, three of the colonies extended their ‘aggressiveness’ by increasing the defender rate with the foraging level. Such ‘super-aggressive’ colonies obviously are able to allocate two separate recruitment pools for foragers and flying defenders.     

Ecological characterization of coral growth anomalies on Porites compressa in Hawai��i

Bulbous skeletal structures with associated aberrant corallites have been abundant on Porites compressa in Kāne‘ohe Bay, O‘ahu, Hawai‘i, for at least the last 19 years. These growth anomalies (GA) appear in the summer in shallow (<3 m) water on some, but not on all colonies. GA-free branches, collected from colonies with GAs, produced GAs when cultured in outdoor flow-thru aquaria. Normal branches, whose tissues were continuous with those of GAs, grew in length much more slowly than normal branches from the same colony that were not connected with a GA, suggesting that there is a translocation of materials from normal tissue to GAs. Small experimental colonies that were either exposed to, or protected from, UV radiation did not differ in their rate of GA formation. GAs had a lower probability of survival than normal branches. This characteristic, in combination with their effect on the growth of normal branches and other reported deficiencies in the tissues of growth anomalies (e.g., reduced or failed reproduction), suggests that GA-bearing colonies of this species have reduced fitness.     

Skeletal development in <Emphasis Type="Italic">Acropora palmata</Emphasis> (Lamarck 1816): a scanning electron microscope (SEM) comparison demonstrating similar mechanisms of skeletal extension in axial versus encrusting growth

Many Acropora palmata colonies consist of an encrusting basal portion and erect branches. Linear growth of the skeleton results in extension along the substrate (encrusting growth), lengthening of branches (axial growth) and thickening of branches and crust (radial growth). Scanning Electron Microscopy is used to compare the mechanisms of skeletal extension between encrusting growth and axial growth. In encrusting growth, the distal margin of the skeleton lacks corallites (which develop about 1 mm from the edge); in contrast, in axial growth, axial corallites along the branch tip form the distal portion of the skeleton. In both locations, the distal margin of the skeleton consists of a lattice-like structure composed of rods that extend from the body of the skeleton and bars that connect these rods. An actively extending skeleton is characterized by sharply pointed rods and partially developed bars. Distal growth of rods (and formation of bars) is effected by the formation of new sclerodermites. Each sclerodermite begins with the deposition of fusiform crystals (that range in length from 1 to 5 μm). These provide a surface for nucleation and growth of spherulitic tufts, clusters of short (<1 μm long) aragonite needles. The needles that are oriented perpendicular to the axis of the skeletal element (rod or bar), and perpendicular to the overlying calicoblastic epithelium, continue extension to appear on the surface of the skeleton as 10–15 μm wide bundles (of needle tips) called fasciculi. However, some crusts that abut competitors for space have a different morphology of skeletal elements (rods and bars). The distal edge of these crusts terminates in blunt coalescing rods, and bars that are fully formed. Absence of fusiform crystals, lack of sharply pointed rods and bars, and full development of sclerodermites characterize a skeletal region that has ceased, perhaps only temporarily, skeletal extension.     

The biology of coral metamorphosis: molecular responses of larvae to inducers of settlement and metamorphosis

Like many other cnidarians, corals undergo metamorphosis from a motile planula larva to a sedentary polyp. In some sea anemones such as Nematostella this process is a smooth transition requiring no extrinsic stimuli, but in many corals it is more complex and is cue-driven. To better understand the molecular events underlying coral metamorphosis, competent larvae were treated with either a natural inducer of settlement (crustose coralline algae chips/extract) or LWamide, which bypasses the settlement phase and drives larvae directly into metamorphosis. Microarrays featuring > 8000 Acropora unigenes were used to follow gene expression changes during the 12 h period after these treatments, and the expression patterns of specific genes, selected on the basis of the array experiments, were investigated by in situ hybridization. Three patterns of expression were common—an aboral pattern restricted to the searching/settlement phase, a second phase of aboral expression corresponding to the beginning of the development of the calicoblastic ectoderm and continuing after metamorphosis, and a later orally-restricted pattern.