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1.
This study investigated the morphology, severity, and distribution of growth anomalies (GAs) in the coral, Montipora capitata, from Wai‘ōpae tide pools, southeast Hawai‘i Island. A macro-image analysis of skeletal microstructure placed GAs into two definable categories; Type A and Type B. Type A GAs had polyp density reduced by 43.05 ± 0.80% (mean ± SE) compared to healthy M. capitata tissue, with many fused and protrusive tuberculae. Type B GAs had no discernable polyps or calices and fused protuberant coenosteum. The prevalence of Type A and Type B GAs among all M. capitata colonies (n = 1,093) in 8 tide pools at Wai‘ōpae was 22.1% (range 2.8–33.7%) and 8.2% (range 0.0–16.9%), respectively. The proportion of colony surface area occupied by GA (relative GA cover) was quantified to assess the severity of this disease among all surveyed colonies. The relative GA cover was significantly greater on colonies larger than 1 m in diameter than smaller colonies and in the central portion of colonies than in the periphery. Furthermore, relative GA cover was negatively related to water motion (R 2 = 0.748, P < 0.01). Developing field diagnostic criteria of M. capitata GA allowed for a detailed epizootiological assessment that determined several cofactors associated with disease severity. Such epizootiological analysis is applicable to future studies of GAs elsewhere.  相似文献   

2.
In this study we provide experimental evidence of transmission of growth anomalies (GAs) between corals. Twenty-four aquaria (16 experimental, 8 containing only apparently healthy corals) were set-up on Negros Island, Philippines, to test for direct-contact and waterborne transmission of GAs. Within seven weeks, two of 16 apparently healthy colonies placed in direct contact with colonies having GAs developed multiple GA lesions whose size and number increased over time. One of 16 apparently healthy colonies in experimental aquaria not touching any diseased colony also developed a GA, exhibiting a single lesion that did not increase in size. Apparently healthy colonies (n=24) in aquaria without a diseased colony remained unchanged.  相似文献   

3.
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.  相似文献   

4.
Burns JH  Takabayashi M 《PloS one》2011,6(12):e28854
Growth anomalies (GAs) affect the coral, Montipora capitata, at Wai'ōpae, southeast Hawai'i Island. Our histopathological analysis of this disease revealed that the GA tissue undergoes changes which compromise anatomical machinery for biological functions such as defense, feeding, digestion, and reproduction. GA tissue exhibited significant reductions in density of ova (66.1-93.7%), symbiotic dinoflagellates (38.8-67.5%), mesenterial filaments (11.2-29.0%), and nematocytes (28.8-46.0%). Hyperplasia of the basal body wall but no abnormal levels of necrosis and algal or fungal invasion was found in GA tissue. Skeletal density along the basal body wall was significantly reduced in GAs compared to healthy or unaffected sections. The reductions in density of the above histological features in GA tissue were collated with disease severity data to quantify the impact of this disease at the colony and population level. Resulting calculations showed this disease reduces the fecundity of M. capitata colonies at Wai'ōpae by 0.7-49.6%, depending on GA severity, and the overall population fecundity by 2.41±0.29%. In sum, GA in this M. capitata population reduces the coral's critical biological functions and increases susceptibility to erosion, clearly defining itself as a disease and an ecological threat.  相似文献   

5.
Growth anomalies (GAs), one of the diseases recently reported for scleractinian corals, are characterized by an abnormal skeletal structure and reduced zooxanthella density. The pathological characteristics of GAs were studied in colonies of Porites australiensis on a reef in Kayo, Okinawa, Japan. Corallites in the GA region lost the skeletal architecture characteristic of P. australiensis, and polyp density had decreased in the GAs due to enlargement of both calices and the coenosteum. The gross productivity of isolated GA samples was lower than in healthy samples and decreased to almost 0 within 11 d after isolation. However, when GA samples were brought into contact with healthy-looking samples from the same colony, they fused and both the GA and healthy regions grew. Healthy samples fused with GA samples grew more slowly than those fused with healthy samples. For in situ GAs surrounded by healthy tissue, tissue death usually started at the center of the GA, probably due to a deficiency in the translocated energy supply from the surrounding tissue. The total area of the GA region and the dead area increased at a rate of 5.3 ± 2.9 cm2 yr-1. These results suggest that GA regions are maintained by energy supplies from surrounding healthy tissues and that GAs may have a negative impact on host corals.  相似文献   

6.

It is critical to determine the methods by which coral colonies regenerate tissue lost to physical injury as they provide the physical structure of coral reef systems. To explore regeneration, circular lesions (12 mm diameter × 3 mm depth) were created in the fall of 2014 on 124 Montastraea cavernosa colonies located in the coastal waters of Grenada and Carriacou (10–12 m depth). Coral regeneration was documented at weekly intervals for 28 days. Repeated measures ANOVA on estimated weekly coral regeneration rates showed that island (p = 0.024) and colony colour (p = 0.024) were the only factors significantly affecting lesion regeneration. Mean rate of lesion closure during the first 28 days was approximately 2.8 mm2 d−1. Four identical circular lesions were created on 30 M. cavernosa colonies (Carriacou, 10–12 m depth) in the fall of 2015. One representative lesion created on each coral colony was re-sampled at each of 14, 21, and 32 or 33 days following injury, and coral tissue was flash-frozen. Tissues from 10 normally pigmented brown colonies were selected for proteomic analysis using tandem mass tags. The initial polyp sample, the day 14, and the final samples were used to quantify the difference in protein abundance as the lesions healed. In the tissue samples 6419 peptides were reliably identified, which corresponded to 906 unique proteins. During the first month of regeneration, 111 proteins were differentially abundant (p < 0.05) on at least one timepoint and of these, 11 were associated with regeneration. An additional 14 proteins were also identified that were differentially abundant (p < 0.05) and were associated with inflammation or antioxidant activity. This work demonstrates, for the first time, the differential abundance of proteins associated with regeneration in a scleractinian coral.

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7.
In addition to experimental studies, computational models provide valuable information about colony development in scleractinian corals. Using our simulation model, we show how environmental factors such as nutrient distribution and light availability affect growth patterns of coral colonies. To compare the simulated coral growth forms with those of real coral colonies, we quantitatively compared our modelling results with coral colonies of the morphologically variable Caribbean coral genus Madracis. Madracis species encompass a relatively large morphological variation in colony morphology and hence represent a suitable genus to compare, for the first time, simulated and real coral growth forms in three dimensions using a quantitative approach. This quantitative analysis of three-dimensional growth forms is based on a number of morphometric parameters (such as branch thickness, branch spacing, etc.). Our results show that simulated coral morphologies share several morphological features with real coral colonies (M. mirabilis, M. decactis, M. formosa and M. carmabi). A significant correlation was found between branch thickness and branch spacing for both real and simulated growth forms. Our present model is able to partly capture the morphological variation in closely related and morphologically variable coral species of the genus Madracis.  相似文献   

8.
We conducted gross and microscopic characterizations of lesions in Cnidaria from Palmyra Atoll, Central Pacific. We found growth anomalies (GA) to be the most commonly encountered lesion. Cases of discoloration and tissue loss were rare. GAs had a focal or multi-focal distribution and were predominantly nodular, exophytic, and umbonate. In scleractinians, the majority of GAs manifested as hyperplasia of the basal body wall (52% of cases), with an associated absence or reduction of polyp structure (mesenteries and filaments, actinopharynx and tentacles), and depletion of zooxanthellae in the gastrodermis of the upper body wall. In the soft corals Sinularia sp. and Lobophytum sp., GAs exclusively manifested as prominent hyperplasia of the coenenchyme with an increased density of solenia. In contrast to scleractinians, soft coral GAs displayed an inflammatory and necrotizing component with marked edema of the mesoglea, accompanied by infiltrates of variably-sized granular amoebocytes. Fungi, algae, sponges, and Crustacea were present in some scleractinian GAs, but absent in soft coral GAs. Fragmentation of tissues was a common finding in Acropora acuminata and Montipora cf. dilatata colonies with tissue loss, although no obvious causative agents were seen. Discoloration in the zoanthid, Palythoa tuberculosa, was found to be the result of necrosis, while in Lobophytum sp. discoloration was the result of zooxanthellar depletion (bleaching). Soft corals with discoloration or tissue loss showed a marked inflammatory response, however no obvious causative organisms were seen. Lesions that appeared similar at the gross level were revealed to be distinct by microscopy, emphasizing the importance of histopathology.  相似文献   

9.
Winter canola (Brassica napus cv Crystal) is an oilseed crop that requires vernalization (chilling treatment) for the induction of stem elongation and flowering. To investigate the role of gibberellins (GAs) in vernalization-induced events, endogenous GA content and the metabolism of [3H]GAs were examined in 10-week vernalized and nonvernalized plants. Shoot tips were harvested 0, 8, and 18 d postvernalization (DPV), and GAs were purified and quantified using 2H2-internal standards and gas chromatography-selected ion monitoring. Concentrations of GA1, GA3, GA8, GA19, and GA20 were 3.1-, 2.3-, 7.8-, 12.0-, and 24.5-fold higher, respectively, in the vernalized plants at the end of the vernalization treatment (0 DPV) relative to the nonvernalized plants. Thermoregulation apparently occurs prior to GA19 biosynthesis, since vernalization elevated the concentration of all of the monitored GAs. [3H]GA20 or [3H]GA1 was applied to the shoot tips of vernalized and nonvernalized plants, and after 24 h, plants were harvested at 6, 12, and 15 DPV. Following high-performance liquid chromatography analyses, vernalized plants showed increased conversion of [3H]GA20 to a [3H]GA1-like metabolite and reduced conversion of [3H]GA1 or [3H]GA20 to polar 3H-metabolites, putative glucosyl conjugates. These results demonstrate that vernalization influences GA content and GA metabolism, with GAs serving as probable regulatory intermediaries between chilling treatment and subsequent stem growth.  相似文献   

10.
Massive colonies of Porites lobata on the barrier reef of Tiahura, Moorea, can be divided into four categories: living colonies, colonies consisting of 50% live coral and 50% dead skeleton, 100% dead coral and colonies which have been reduced to a basal plate. Replicate samples of each of these colony types were collected in the same vicinity of the barrier reef during October 1987. The macroborers were extracted, identified, counted and their volumes determined by displacement. Kruskal-Wallis tests showed that three different boring communities occur within these four categories of Porites colonies. Live colonies are characterised by only 3 species, the bivalve, Lithophaga laevigata; the vermetid Dendropoma maximun; and the non boring serpulid polychaete Spirobranchus. The completely dead colonies contain up to 17 boring species, with five to six individuals per 100 cm. Sipunculans are the dominant bioeroders with the most abundant species being Aspidosiphon elegans, sp A and sp B. Colonies of Porites which have been reduced to basal plates contain up to 18 boring species of which the bivalve Lithophaga hanleyana and the sipunculan Aspidosiphon sp. B are the most abundant.The cumulative volume of CaCO3 lost by boring activity increases from 0.1 cm3 per 100 cm3 in a completely dead Porites colony to 1.4 cm3 per 100 cm3 in the residual basal plates of Porites. These can be extrapolated to minimum losses of 14.2 kg m-3. We suggest that rates of boring increase with the time which has elapsed since the death of the colony and the dominant agents of boring also change with increasing age of the coral structure. There are significant additional losses of 5.25 kg m-3 CaCO3 caused by grazing echinoids and scarids.  相似文献   

11.
In G2 peas (Pisum sativum L.) apical senescence occurs only in long days (LD), and indeterminate growth is associated with elevated gibberellin (GA) levels in the shoot in short days (SD). Metabolism of GA12 aldehyde was investigated by feeding shoots grown in SD or LD with [14C]GA12 aldehyde through the cut end of the stem for 0.5 to 6 hours in the light and analyzing the tissue extract by high performance liquid chromatography. More radioactive products were detected than can be accounted for by the two GA metabolic pathways previously known to be present in peas. Three of the major products appear to be GA conjugates, but an additional pathway(s) of GA metabolism may be present. The levels of putative C20 GAs, [14C]GA53, [14C]GA44, [14C]GA19, and/or [14C] GA17, were all elevated in SD as compared to LD. Putative [14C]GA, was slightly higher in LD than in SD. Putative [14C]GA53 was a major metabolite after 30 minutes of treatment in SD but had declined after longer treatment times to be replaced by elevated levels of putative [14C] GA44 and [14C]GA19/17. Metabolism of GA20 was slow in both photoperiods. Although GA20 and GA19 are the major endogenous GAs as determined by gas chromatography-mass spectrometry, putative [14C]GA20 and [14C]GA19 were never major products of [14C]GA12 aldehyde metabolism. Thus, photoperiod acts in G2 peas to change the rate of GA53 production from GA12 aldehyde, with the levels of the subsequent GAs on the 13-OH pathway being determined by the amount of GA53 being produced.  相似文献   

12.
The application of gibberellin A4/7 (GA4/7) to the stem of previous-year (1-year-old) terminal shoots of Scots pine (Pinus sylvestris) seedlings has been observed to stimulate cambial growth locally, as well as at a distance in the distal current-year terminal shoot, but the distribution and metabolic fate of the applied GA4/7, as well as the pathway of endogenous GA biosynthesis in this species, has not been investigated. As a first step, we analysed for endogenous GAs and monitored the transport and metabolism of labelled GAs 4, 9 and 20. Endogenous GAs from the elongating current-year terminal shoot of 2-year-old seedlings were purified by column chromatography and high-performance liquid chromatography and analysed by combined gas chromatography-mass spectrometry (GC-MS). GAs 1, 3, 4, 9, 12 and 20 were identified in the stem, and GAs 1, 3 and 4 in the needles, by full-scan mass spectrometry (GAs 1, 3, 4, 9 and 12) or selected-ion monitoring (GA20) and Kovats retention index. Tritiated and deuterated GA4, GA9 or GA20 were applied around the circumference at the midpoint of the previous-year terminal shoot, and metabolites were extracted from the elongating current-year terminal shoot, the application point, and the 1-year-old needles and the cambial region above and below the application point. After purification, detection by liquid scintillation spectrometry and analysis by GC-MS, it was evident that, for each applied GA, unmetabolised [2H2]GA and [3H]radioactivity were present in every seedling part analysed. Most of the radioactivity was retained at the application point when [3H]GA9 and [3H]GA20 were applied, whereas the largest percentage of radioactivity derived from [3H]GA4 was recovered in the current-year terminal shoot. It was also found that [2H2]GA9 was converted to [2H2]GA20 and to both [2H2]GA4 and [2H2]GA1, [2H2]GA4 was metabolised to [2H2]GA1, and [2H2]GA20 was converted to [2H2]GA29. The data indicate that for Pinus sylvestris shoots (1) GAs applied laterally to the outside of the vascular system of previous-year shoots not only are absorbed and translocated extensively throughout the previous-year and current-year shoots, but also are readily metabolised, (2) the GA metabolic pathways found are closely related to the endogenous GAs identified, and (3) GA9 metabolism follows two distinctly different routes: in one, GA9 is converted to GA1 through GA4, and in the other it is converted to GA20, which is then metabolised to GA29. The results suggest that the late 13-hydroxylation pathway is an important route for GA biosynthesis in shoots of Pinus sylvestris, and that the stimulation of cambial growth in Scots pine by exogenous GA4/7 may be due to its conversion to GA1, rather than to it being active per se.  相似文献   

13.
This study describes the distribution and abundance patterns of the associate fauna on the living surface of the corals Siderastrea stellata Verril, 1868 and Mussismilia hispida (Verril 1902) using a non-destructive method, on the northern coast of Rio de Janeiro State. For each coral species, infestation density and proportions of infested colonies, colonies attached and unattached to the substrate were estimated. A total of 474 colonies of S. stellata and 452 colonies of M. hispida were examined. The barnacle Ceratoconcha floridana (Pilsbry, 1931) was the dominant coral associate found, followed by gall-crabs of the family Cryptochiridae Paulson, 1875 and the bivalve Lithophaga bisulcata (d’Orbigny, 1842). Both coral species presented similar patterns of infestation dominance. S. stellata colonies were more commonly infested and showed a greater mean infestation density of 0.62 ind/cm2 at Armação dos Búzios, whereas M. hispida colonies had infestation densities of only 0.20 ind/cm2. Infestation density does not appear to impact negatively on corals of Armação dos Búzios. A clear negative relationship between the number of associates in the coral colony and coral size was found. Evidently abundance and frequency of occurrence of associated fauna is highly related to coral community structure and composition and the results highlight the importance of local scale studies.  相似文献   

14.
Bastidas  C.  Fabricius  K. E.  Willis  B. L. 《Hydrobiologia》2004,530(1-3):433-441
We evaluated the role that demography may play in the formation of local aggregations of Sinularia flexibilis (Quoy & Gaimard, 1833), a soft coral that commonly dominates inshore coral reefs of the Great Barrier Reef (GBR), Australia. Two populations on inshore reefs of the Palm Islands were censused once a year for 3 years, starting 10 mo after the extensive bleaching mortality in early 1998. Larger colonies became more prevalent over time; mean colony size increasing by 35%, from 276 cm2 in 1998 to 373 cm2 in 2000. Growth rates were size dependent, with smaller colonies growing proportionally faster than larger colonies. Change in size relative to initial size indicated an expected mean annual growth of 128 cm2 for a 50-cm2 colony. Zero growth was predicted at 532±21cm2, with colonies larger than this likely to undergo fission or shrink. Forty-three percent of colonies were undergoing fission at any time at both localities. Most new colonies were produced by fission (70%, n=285), with the remainder produced by the recruitment of sexually produced larvae (19%) or by colony translocation (11%). The sexual and asexual recruitment rates were 0.24 and 1.0 recruits m- 2 year−1, respectively. Opportunistic recruitment and rapid growth following disturbances are commonly assumed to be the mechanisms leading soft corals to dominate locally. In this study, these mechanisms operated more slowly than expected, with no net change in population size.  相似文献   

15.
Localized bleaching (a discrete white area on the coral) was observed in one of the main framework-building corals in Hawaii, Porites compressa. This study aimed to determine the degree of virulence of the lesion. We investigated the whole-colony effects by following disease progression through time and examining the effect of localized bleaching on coral fecundity. After two months, 35 of 42 (83.3%) individually tagged colonies affected by localized bleaching showed tissue loss and partial colony mortality. Histological slides of healthy P. compressa and samples from colonies showing signs of localized bleaching were compared showing that affected colonies had a significant reduction (almost 50%) in gamete development, egg numbers, and egg size in the affected tissue. The observed localized bleaching results in both partial colony mortality and a reduced number of gametes and was termed Porites Bleaching with Tissue Loss (PBTL).  相似文献   

16.
The sediment rejection abilities of colonies of Montastrea cavernosa (Linnaeus) were monitored in a series of field and laboratory experiments. Under conditions of natural sediment deposition, colonies were capable of removing virtually all sediment from their surfaces. Equivalent sections of dead coral skeleton accumulated as much as 345 mg sediment · 25 cm?2 · day?1. Laboratory experiments show the sediment rejection process to consist of passive and active phases. Passive movement of sediment off colonies occurs at the time of initial sediment impact. Fine grain size, tall polyps, and convex colonies all enhance passive removal. Active removal involves the behavioral responses of the colony and is independent of sediment type or colony morphology. The presence of appropriate morphologies can increase the proportion of sediment passively removed and reduce the amount of work a colony must perform to keep its surface clear of sediment.  相似文献   

17.
We assessed the distribution and prevalence of growth anomalies (GAs) in Acropora from French Frigate Shoals (Hawaii, USA), Johnston Atoll and Tutuila (American Samoa), developed a nomenclature for gross morphology, characterized GAs at the cellular level and obtained preliminary indices of their spatial patterns and progression within coral colonies. Acropora GAs were found in all 3 regions, but the distribution, variety and prevalence of Acropora GAs was highest in American Samoa. GAs were grouped into 7 gross morphologies (exophytic, bosselated, crateriform, nodular, vermiform, fimbriate or annular). On histology, GAs consisted of hyperplastic basal body wall (calicodermis, mesoglea and gastrodermis apposed to skeleton) with 3 distinct patterns of necrosis. There was no evidence of anaplasia or mitotic figures (common but not necessarily required morphologic indicators of neoplasia). Compared to normal tissues, GAs had significantly fewer polyps, zooxanthellae within the gastrodermis of the coenenchyme, mesenterial filaments and gonads but significantly more necrosis. On 2 colonies with GAs monitored at 2 points over 11 mo, numbers of GAs per colony increased from 0.9 to 3 times the original number seen, and significant clustering of GAs occurred within colonies. The evidence of GAs being true neoplasias (tumors) is mixed, so a cautionary approach is urged in use of morphologic terminology.  相似文献   

18.
The octocoral Heliopora coerulea is a gonochoric surface brooder. Although the species is common on shallow Indo-Pacific coral reefs, information on its reproductive biology is limited and spawning timings have only been reported from four locations. We report the first observations of surface brooding in H. coerulea on a sedimented equatorial reef. In April 2014, 46 and 26 brooding colonies were recorded, respectively, from reefs fringing the islands of Kusu and Lazarus in Singapore. The brooded coral larvae were tightly bound to inflated coral polyps, forming a dense white mat which blanketed the brown coral tissue. The inflated polyps also protruded from layers of sediment, which accumulated in colony crevices, such that the brooded larvae were elevated above the settled sediment. Our observations supplement existing knowledge on the reproductive timing of H. coerulea and highlight the adaptations that improve survival of this species on sediment-impacted reefs.  相似文献   

19.
To examine algal (= zooxanthellae) regulation and control, and the factors determining algal densities in hard corals, the zooxanthellae mitotic index and release rates were regularly determined in branch tips from a colony of a staghorn coral, Acropora formosa, recovering from a coral ''bleaching'' event (the stress-related dissociation of the coral–algal symbiosis). Mathematical models based upon density-dependent decreases in the algal division frequency and increases in algal release rates during the post-bleaching recovery period accurately predict the observed recovery period (ca. 20 weeks). The models suggest that (i) the colony recovered its algal population from the division of the remaining zooxanthellae, and (ii) the continual loss of zooxanthellae significantly slowed the recovery of the coral. Possible reasons for the ''paradoxical'' loss of healthy zooxanthellae from the bleached coral are discussed in terms of endodermal processes occurring in the recovering coral and the redistribution of newly formed zooxanthellae to aposymbiotic host cells. At a steady-state algal density of 2.1 x 106 zooxanthellae cm-2 at the end of the recovery period, the zooxanthellae would have to form a double layer of cells in the coral tissues, consistent with microscopic observations. Neighbouring colonies of A. formosa with inherently higher algal densities possess proportionately smaller zooxanthellae. Results suggest that space availability and the size of the algal symbionts determines the algal densities in the coral colonies. The large increases in the algal densities reported in corals exposed to elevated nutrient concentrations (i.e between a two- and five-fold increase in the algal standing stock) are not consistent with this theory. We suggest that increases of this magnitude are a product of the experimental conditions: reasons for this statement are discussed. We propose that the stability of the coral–algal symbiosis under non-stress conditions, and the constancy of zooxanthellae densities in corals reported across growth form, depth and geographic range, are related to space availability limiting algal densities. However, at these densities, zooxanthellae have attributes consistent with nutrient limitation.  相似文献   

20.
‘White syndrome’ is considered to be the most prevalent coral disease on the Great Barrier Reef, characterised by rapid rates of lesion progression and high levels of colony mortality. This study investigated the production and translocation of photoassimilates towards white syndrome lesions (WSLs) and artificially inflicted lesions in healthy and diseased colonies of tabular Acropora spp. to determine the intra-colonial response to white syndrome using 14C labelling. Translocation of 14C labelled photoassimilates was preferentially orientated away from active WSLs, with minimal 14C activity observed in the lesion borders, whilst artificial lesions (ALs) created directly opposite WSL borders showed significantly higher 14C activity, suggesting active translocation of photoassimilates for tissue regeneration. Transport of photoassimilates in healthy coral colonies was preferentially oriented towards ALs with a higher perimeter–area ratio, although translocation towards WSL boundaries was minimal even though the lesion perimeter was often the width of the colony (>200 cm). We suggest that the preferential orientation of photoassimilates away from WSLs may represent a deliberate strategy by the colony to induce a ‘shutdown reaction’ in order to preserve intra-colonial resources within areas of the colony that are more likely to survive and recover.  相似文献   

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