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1.
A study of new Halimeda material available from the International Indian Ocean Expedition, and of type and other specimens not previously available to me, further indicated that Halimeda opuntia is a composite of species rather than a single species. The characteristics of some of its infraspecific taxa differ sufficiently from typical H. opuntia that I have elevated them to species. Extended diagnoses of three, H. hederacea, H. minima, and H. distorta, are given in this paper. Some characteristics new in Halimeda taxonomy are introduced, and distinguishing macroscopic and microscopic characteristics for the new species are discussed. Available ecological data are included.  相似文献   

2.
A new species, Halimeda hummii, and a new variety of Halimeda cryptica Colinvaux and Graham both originally collected from the edge of the continental shelf on the southwest coast of Puerto Rico are newly described. The new species is irregular in its segment morphology and is the smallest species of Halimeda presently known. Halimeda cryptica var. acerifolia from deep water possesses distinctive segments resembling maple leaves. Halimeda copiosa Goreau and Graham and typical H. cryptica are also collected in deep water and are newly recorded from Puerto Rico.  相似文献   

3.
4.
The ultrastructure of 4 species of the calcareous, siphonaceous alga Halimeda (H. cylindracea Decaisne, H. discoidea Decaisne, H. macroloba Decaisne and H. tuna (Ellis & Solander) Lamour) has been studied, and the observed changes during growth and development are related to changes in the degree of calcification. A distinct gradient in the types and quantities of cell organelles exists in a growing apical filament. As these filaments grow, branch, and eventually develop into a mature segment, changes in the organization of organelles such as mitochondria and chloroplasts are observed. Calcification begins when the chloroplasts reach structural maturity and when the peripheral utricles adhere (fuse). This adhesion of the peripheral utricles isolates the intercellular space (ICS) in which calcification occurs from the external seawater. Calcification begins in the outermost (pilose) cell wall layer of the walls facing into the ICS. The cell walls at the thallus exterior undergo extensive changes after utricular fusion; the pilose layer is lost, the cuticles of adjacent utricles fuse forming a ridge at their junction, and multiple cuticles are formed. The aragonite (CaCO3) crystals which are initially precipitated within the pilose wall layer, rapidly increase in size and number, eventually filling much of the ICS. Only the initial nucleation of aragonite is associated with the pilose wall layer, the later precipitation of aragonite is totally independent of the pilose layer. In older segments secondary deposition of CaCO3 also occurs around existing aragonite needles.  相似文献   

5.
The capitular filaments of Penicillus and Rhipocephalus consist of an inner tube containing the cytoplasm and an outer calcified sheath. The sheath originates at the cell wall and differentiates into several layers which form the outer filament wall. CaCO3 is deposited between organic layers within the sheath and is not in direct contact with the seawater. Pores within the sheath, usually uncalcified, may facilitate exchange of gases and solutes. The cytoplasm is characterized by vacuolar inclusions of calcium oxalate needles 50–150 μm long. A closed cortical surface is lacking. Udotea cyathiformis Dec. and U. conglutinata (Ellis & Sol.) Lam. are similar to Penicillus and Rhipocephalus, in addition showing some CaCO3 between filaments (ICS-calcification). Udotea flabellum (Ellis & Sol.) Lam. is different as the filaments are profusely branched giving rise to a fully developed cortical surface. Pores and vacuolar calcium oxalate inclusions are absent. CaCO3 deposition occurs within cortical filaments in between layers of the filament wall and subcortically in intercellular spares (ICS). Cortex calcification shows primary and secondary deposits bearing some resemblance to sheath calcification and to coralline red algae. In Rhipocephalus phoenix (Ellis & Sol.) Kütz., Penicillus pyriformis A. &E. Gepp, U. cyathiformis and U. conglutinata CaCO3 is precipitated intracellularly within the sheath, in contrast to Halimeda and Cymopolia where it is deposited extracellularly in between filaments. U. flabellum takes an intermediate position showing both intra- and intercellular calcification. The sheath compartment volume is between 12.5 and 7500 μm3and 5–3 orders of magnitude smaller than the ICS-compartment. Compartment size and location of CaCO3may bear on calcification mechanisms. One condition for such a mechanism may be restricted exchange of solutes (CO2, CO32-, HCO3-, O2, Co2+). Codiaceae; filament ultrastructure; Penicillus; Rhipocephalus; Udotea  相似文献   

6.
The rate of calcification in the scleractinian coral Galaxea fascicularis was followed during the daytime using 45Ca tracer. The coral began the day with a low calcification rate, which increased over time to a maximum in the afternoon. Since the experiments were carried out under a fixed light intensity, these results suggest that an intrinsic rhythm exists in the coral such that the calcification rate is regulated during the daytime. When corals were incubated for an extended period in the dark, the calcification rate was constant for the first 4 h of incubation and then declined, until after one day of dark incubation, calcification ceased, possibly as a result of the depletion of coral energy reserves. The addition of glucose and Artemia reduced the dark calcification rate for the short duration of the experiment, indicating an expenditure of oxygen in respiration. Artificial hypoxia reduced the rate of dark calcification to about 25% compared to aerated coral samples. It is suggested that G. fascicularis obtains its oxygen needs from the surrounding seawater during the nighttime, whereas during the day time the coral exports oxygen to the seawater.  相似文献   

7.
Larkum AW  Salih A  Kühl M 《PloS one》2011,6(7):e20841

Background

The calcifying siphonalean green alga, Halimeda macroloba is abundant on coral reefs and is important in the production of calcium carbonate sediments. The process by which new green segments are formed over-night is revealed here for the first time.

Methodology/Principal Findings

Growth of new segments was visualised by epifluorescence and confocal microscopy and by pulse amplitude modulation (PAM) fluorimetry. Apical colourless proto-segments were initiated on day 1, and formed a loose network of non-calcified, non-septate filaments, containing no chloroplasts. Rapid greening was initiated at dusk by i) the mass movement of chloroplasts into these filaments from the parent segment and ii) the growth of new filaments containing chloroplasts. Greening was usually complete in 3–5 h and certainly before dawn on day 2 when the first signs of calcification were apparent. Mass chloroplast movement took place at a rate of ∼0.65 µm/s. Photosynthetic yield and rate remained low for a period of 1 to several hours, indicating that the chloroplasts were made de novo. Use of the inhibitors colchicine and cytochalasin d indicated that the movement process is dependent on both microtubules and microfilaments.

Significance

This unusual process involves the mass movement of chloroplasts at a high rate into new segments during the night and rapid calcification on the following day and may be an adaptation to minimise the impact of herbivorous activity.  相似文献   

8.
Abstract

Halimeda has been found particularly suitable for studies of long‐distance chloroplast migration by virtue of its coenocytic structure and calcium carbonate skeleton. A circadian rhythm of chloroplast migration in Halimeda distorta was monitored by videography of segment surface pigmentation. In normal 12 h light/12 h dark treatments synchronised with dawn and dusk, the segments were green all day, began to become pale immediately the light was turned off, and then remained almost white for most of the night until beginning to re‐green a few hours before dawn. As a result of that, they were already quite green by the time the light was turned on. In continuous darkness a similar cycle, albeit with reducing amplitude and a period of about 23 hours, was maintained for at least 7 days. However, this cycle differed significantly from the normal one in that the segments did not remain green after the light was not switched on at dawn, but rather began to pale immediately thereafter. Conversely, in continuous light the segments did not become pale at any time. Thus, the rhythmical re‐emergence of the chloroplasts before dawn and their subsequent withdrawal appears to be controlled by an endogenous rhythm which is independent of light. However, light does completely, but reversibly, inhibit the chloroplast withdrawal component of the cycle. This behaviour of the chloroplasts in Halimeda is very similar to that in the related alga, Caulerpa, but it is quite different from that in another extensively Studied but unrelated siphonous green alga, Acetabularia, in which the circadian rhythm of chloroplast migration is maintained in continuous light.  相似文献   

9.
McNicholl  C.  Koch  M. S.  Swarzenski  P. W.  Oberhaensli  F. R.  Taylor  A.  Batista  M. Gómez  Metian  M. 《Coral reefs (Online)》2020,39(6):1635-1647

Net calcification rates for coral reef and other calcifiers have been shown to decline as ocean acidification (OA) occurs. However, the role of calcium carbonate dissolution in lowering net calcification rates is unclear. The objective of this study was to distinguish OA effects on calcification and dissolution rates in dominant calcifying macroalgae of the Florida Reef Tract, including two rhodophytes (Neogoniolithon strictum, Jania adhaerens) and two chlorophytes (Halimeda scabra, Udotea luna). Two experiments were conducted: (1) to assess the difference in gross (45Ca uptake) versus net (total alkalinity anomaly) calcification rates in the light/dark and (2) to determine dark dissolution (45CaCO3), using pH levels predicted for the year 2100 and ambient pH. At low pH in the light, all species maintained gross calcification rates and most sustained net calcification rates relative to controls. Net calcification rates in the dark were ~84% lower than in the light. In contrast to the light, all species had lower net calcification rates in the dark at low pH with chlorophytes exhibiting net dissolution. These data are supported by the relationship (R2 = 0.82) between increasing total alkalinity and loss of 45Ca from pre-labelled 45CaCO3 thalli at low pH in the dark. Dark dissolution of 45CaCO3-labelled thalli was ~18% higher in chlorophytes than rhodophytes at ambient pH, and ~ twofold higher at low pH. Only Udotea, which exhibited dissolution in the light, also had lower daily calcification rates integrated over 24 h. Thus, if tropical macroalgae can maintain high calcification rates in the light, lower net calcification rates in the dark from dissolution may not compromise daily calcification rates. However, if organismal dissolution in the dark is additive to sedimentary carbonate losses, reef dissolution may be amplified under OA and contribute to erosion of the Florida Reef Tract and other reefs that exhibit net dissolution.

  相似文献   

10.
Halimeda opuntia is a cosmopolitan marine calcifying green alga in shallow tropical marine environments. Besides Halimeda’s contribution to a diverse habitat, the alga is an important sediment producer. Fallen calcareous segments of Halimeda spp. are a major component of carbonate sediments in many tropical settings and play an important role in reef framework development and carbonate platform buildup. Consequently the calcification of H. opuntia accounts for large portions of the carbonate budget in tropical shallow marine ecosystems. Earlier studies investigating the calcification processes of Halimeda spp. have tended to focus on the microstructure or the physiology of the alga, thus overlooking the interaction of physiological and abiotic processes behind the formation of the skeleton. By analyzing microstructural skeletal features of Halimeda segments with the aid of scanning electron microscopy and relating their occurrence to known physiological processes, we have been able to identify the initiation of calcification within an organic matrix and demonstrate that biologically induced cementation is an important process in calcification. For the first time, we propose a model for the calcification of Halimeda spp. that considers both the alga’s physiology and the carbon chemistry of the seawater with respect to the development of different skeletal features. The presence of an organic matrix and earlier detected external carbonic anhydrase activity suggest that Halimeda spp. exhibit biotic precipitation of calcium carbonate, as many other species of marine organisms do. On the other hand, it is the formation of micro-anhedral carbonate through the alga’s metabolism that leads to a cementation of living segments. Precisely, this process allows H. opuntia to contribute substantial amounts of carbonate sediments to tropical shallow seas.  相似文献   

11.
The mineral component of the marine green algal genus Halimeda is the orthorhombic calcium carbonate (aragonite); its presence appears to be a generic characteristic. Tydemania expeditionis also precipitates aragonite in contradistinction to species of the red alga Corallina wliich precipitate calcium carbonate of the hexagonal form (calcite). The analyses are based on x-ray diffraction methods. Although other inorganic substances arc present, the amounts are minor and probably represent contaminants. Specimens that are to be studied for mineral components should not be stored in formalin.  相似文献   

12.
The calcified green algal genus Halimeda is one of the most ecologically important but morphologically diverse seaweeds in sub-tropical and tropical waters. Because of its high morphological plasticity, the identification of Halimeda species based on morphological characters is challenging without the assistance of molecular analysis. To date, the species diversity of Halimeda in Taiwan and its overseas territories has not been investigated with the assistance of DNA sequencing, and this taxonomic knowledge gap should be filled. The present study initiates a systematic examination of the species diversity and distribution of Halimeda in Taiwan, Spratly Island, and Dongsha Atoll in the South China Sea, using DNA sequence data (plastid tufA gene and rbcL) and morphological data. Our DNA analyses revealed the presence of 10 Halimeda species (Halimeda borneensis, Halimeda cylindracea, Halimeda discoidea, Halimeda distorta, Halimeda macroloba, Halimeda minima, Halimeda opuntia, Halimeda renschii, Halimeda taiwanensis sp. nov., and Halimeda velasquezii) in the waters around Taiwan, Spratly Island, and Dongsha Atoll. The majority of the species could be readily distinguished by their morphological and anatomical characters. The proposed new species, H. taiwanensis, was differentiated not only by our algorithmic species delimitation analyses (statistical parsimony network analysis and automated barcode gap discovery), but also by its morphological features. The proposed new species differs from two externally resembled species, H. cuneata and H. discoidea, in having an undulated segment margin, the complete fusion of medullary siphons at the node, the lack of segment stalk, and the presence of a large primary utricle. Here, we present the up-to-date taxonomic account, molecular diversity, and geographical distribution of Halimeda spp. in Taiwan and associated areas of the South China Sea. Environmental factors that might drive the occurrence and latitudinal distribution of the species are also discussed.  相似文献   

13.
14.
The effects of a number of metabolic inhibitors on calcificationand photosynthesis in Halimeda tuna, H. discoidea, and H. macrolobaare described. The inhibitors used are CCCP, DNP, DCMU, azide,cyanide, chloramphenicol, cycloheximide, and Diamox. The effectsof these inhibitors, although complex, are consistent with ourmodel of calcification in Halimeda. Inhibition of photosyntheticCO2 uptake inhibits calcification as does stimulation of respiratoryCO2 evolution (i.e. uncoupling). There is also indirect evidencefor the presence of a possible light stimulated H+ efflux whichinhibits calcification. The observed calcification rate is thereforethe result of a number of factors which affect the concentrationof COand the pH in the intercellular space of the Halimedathallus. The results obtained with the carbonic anhydrase inhibitor Diamoxprovide further evidence for the effective separation of theintercellular space from the external medium by the appressedperipheral utricles.  相似文献   

15.
Estimates for the production of calcium carbonate by Halimeda spp. have been based on limited measures in small areas or over short periods, subsequently extrapolated over larger temporal and spatial scales. The accuracy of these extrapolations depends on the variations in time and space of the parameters used for the derivations of the production, which were evaluated in the present study for Halimeda incrassata (J. Ellis) J. V. Lamour. in the Puerto Morelos reef lagoon, Mexican Caribbean. Growth, biomass, and CaCO3 content of the branches of the thalli were determined at 1–3 monthly intervals from November 1997 until June 1999, using the marking technique with the coloring agent Alizarin Red‐S. Biomass exhibited great variability (V=25.4%), and fluctuations were closely related to changes in thallus density. Growth and CaCO3‐content changes were related to the solar cycle, having coefficients of 15.4% and 2.5%, respectively. Additionally, calcified biomasses of the thalli were determined over a large spatial (31 stations in ~7 km2 area) and time (6 y) scale. Spatial variability in calcified biomass was 59%, and the coefficient of variation attained its highest value (69%) for the samples collected over a 6 y period, from 1990 to 1996 (except 1995). Based on overall average values, branches of H. incrassata in the study area had a mean turnover of 30 d, with an annual production of 815 g CaCO3 per square meter.  相似文献   

16.
The results of a study of nutrient enrichment with nitrogen (N) and phosphorus (P) on productivity and calcification of fleshy and calcareous algae are reported in this study. Plants were collected from a nearshore eutrophic site in the Florida Keys (USA) and experimentally pulsed during the night with combinations of N and P. After several days of pulsing (7–10 days), net productivity, calcification, and alkaline phosphatase activity (APA), were measured. Productivity of fleshy algae were frequently enhanced by N, P, and N+P, during both summer and winter. Phosphorus limited the productivity of Hydroclathrus clathratus during winter and Ulva spp. during summer, whereas nitrogen limited the productivity of Laurencia intricata during both seasons. During summer, Dictyota cervicornis productivity was not enhanced by any nutrient enrichment. Nitrogen limited the productivity of the three calcareous species Penicillus capitatus, Penicillus dumetosus and Halimeda opuntia during winter and that of H. opuntia during summer. Neither N nor P enrichment increased calcification of calcareous species, and P enrichment greatly inhibited calcification of P. dumetosus during winter. Nutrient enrichment enhanced the productivity of the fleshy species to a greater extent than that of calcareous algae. The seawater DIN:SRP molar ratio was low at our eutrophic study site (molar ratio average of 3:1 during winter and 9:1 during summer) compared to more oligotrophic sites in the Florida Keys, suggesting that in carbonate-rich environments, eutrophication shifts nutrient regulation of productivity from P to N. APA activities of fleshy macroalage were higher than calcareous algae, and rates of all macro algae were 2- to 7-fold higher in summer compared to winter. Productivity was also about 3-fold higher in fleshy compared to calcareous species and about 2-fold higher in summer compared to winter. These results suggest that nutrient enrichment enhances productivity of fleshy algae to a greater extent than that of calcareous algae. Thus, overgrowth of calcareous algae by more opportunistic fleshy forms could reduce carbonate accretion in tropical coastlines experiencing increased eutrophication.  相似文献   

17.
With microsensors, we measured the steady‐state microprofiles of O2, pH and Ca2+ on the topside of young segments of Halimeda discoidea, as well as the surface dynamics upon light–dark shifts. The effect of several inhibitors was studied. The steady‐state measurements showed that under high light intensity, calcium and protons were taken up, while O2 was produced. In the dark, O2 was consumed, the pH decreased to below seawater level and Ca2+ uptake was reduced to 50%. At low light intensity (12 mmol photons m‐2 s‐1), Ca2+ efflux was observed. Upon light–dark shifts, a complicated pattern of both the pH and calcium surface dynamics was observed. Illumination caused an initial pH decrease, followed by a gradual pH increase: this indicated that the surface pH of H. discoidea is determined by more than one light‐induced process. When photosynthesis was inhibited by dichlorophenyl dimethyl urea (DCMU), a strong acidification was observed upon illumination. The nature and physiological function of this putative pump is not known. The calcium dynamics followed all pH dynamics closely, both in the presence and absence of DCMU. The Ca‐channel blockers verapamil and nifedipine had no effect on the Ca2+ dynamics and steady‐state profiles. Thus, in H. discoidea, calcification is not regulated by the alga, but is a consequence of pH increase during photosynthesis. Acetazolamide had no effect on photosynthesis, whereas ethoxyzolamide inhibited photosynthesis at higher light intensities. Therefore, all carbonic anhydrase activity is intracellular. Carbonic anhydrase is required to alleviate the CO2 limitation. Calcification cannot supply sufficient protons and CO2 to sustain photosynthesis.  相似文献   

18.
The response of respiration, photosynthesis, and calcification to elevated pCO2 and temperature was investigated in isolation and in combination in the Mediterranean crustose coralline alga Lithophyllum cabiochae. Algae were maintained in aquaria during 1 year at near‐ambient conditions of irradiance, at ambient or elevated temperature (+3°C), and at ambient (ca. 400 μatm) or elevated pCO2 (ca. 700 μatm). Respiration, photosynthesis, and net calcification showed a strong seasonal pattern following the seasonal variations of temperature and irradiance, with higher rates in summer than in winter. Respiration was unaffected by pCO2 but showed a general trend of increase at elevated temperature at all seasons, except in summer under elevated pCO2. Conversely, photosynthesis was strongly affected by pCO2 with a decline under elevated pCO2 in summer, autumn, and winter. In particular, photosynthetic efficiency was reduced under elevated pCO2. Net calcification showed different responses depending on the season. In summer, net calcification increased with rising temperature under ambient pCO2 but decreased with rising temperature under elevated pCO2. Surprisingly, the highest rates in summer were found under elevated pCO2 and ambient temperature. In autumn, winter, and spring, net calcification exhibited a positive or no response at elevated temperature but was unaffected by pCO2. The rate of calcification of L. cabiochae was thus maintained or even enhanced under increased pCO2. However, there is likely a trade‐off with other physiological processes. For example, photosynthesis declines in response to increased pCO2 under ambient irradiance. The present study reports only on the physiological response of healthy specimens to ocean warming and acidification, however, these environmental changes may affect the vulnerability of coralline algae to other stresses such as pathogens and necroses that can cause major dissolution, which would have critical consequence for the sustainability of coralligenous habitats and the budgets of carbon and calcium carbonate in coastal Mediterranean ecosystems.  相似文献   

19.
The photosynthetic performance, pigmentation, and growth of a Halimeda community were studied over a depth gradient on Conch Reef, Florida Keys, USA during summer–fall periods of 5 consecutive years. The physiology and growth of H. tuna (Ellis & Solander) Lamouroux and H. opuntia (L.) Lamouroux on this algal dominated reef were highly variable. Maximum rate of net photosynthesis (Pmax), respiration rate, and quantum efficiency (α) did not differ between populations of either species at 7 versus 21 m, even though the 21‐m site received a 66% lower photon flux density (PFD). Physiological parameters, as well as levels of photosynthetic pigments, varied temporally. Pmax, saturation irradiance, compensation irradiance, and growth were greatest in summer months, whereas α, chl a, chl b, and carotenoid concentrations were elevated each fall. Halimeda tuna growth rates were higher at 7 m compared with 21 m for only two of five growth trials. This may have arisen from variability in light and nutrient availability. Individuals growing at 7 m received a 29% greater PFD in August 2001 than in 1999. In August 1999 and 2001 seawater temperatures were uniform over the 14‐m gradient, whereas in August 2000 cold water regularly intruded upon the 21‐m but not the 7‐m site. These results illustrate the potentially dynamic relationship between nutrients, irradiance, and algal productivity. This suggests the necessity of long‐term monitoring over spatial and temporal gradients to accurately characterize factors that impact productivity.  相似文献   

20.
Previous studies have demonstrated that coral and algal calcification is tightly regulated by the calcium carbonate saturation state of seawater. This parameter is likely to decrease in response to the increase of dissolved CO2 resulting from the global increase of the partial pressure of atmospheric CO2. We have investigated the response of a coral reef community dominated by scleractinian corals, but also including other calcifying organisms such as calcareous algae, crustaceans, gastropods and echinoderms, and kept in an open‐top mesocosm. Seawater pCO2 was modified by manipulating the pCO2 of air used to bubble the mesocosm. The aragonite saturation state (Ωarag) of the seawater in the mesocosm varied between 1.3 and 5.4. Community calcification decreased as a function of increasing pCO2 and decreasing Ωarag. This result is in agreement with previous data collected on scleractinian corals, coralline algae and in a reef mesocosm, even though some of these studies did not manipulate CO2 directly. Our data suggest that the rate of calcification during the last glacial maximum might have been 114% of the preindustrial rate. Moreover, using the average emission scenario (IS92a) of the Intergovernmental Panel on Climate Change, we predict that the calcification rate of scleractinian‐dominated communities may decrease by 21% between the pre‐industrial period (year 1880) and the time at which pCO2 will double (year 2065).  相似文献   

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