Mangroves are woody plants that grow at the interface between land and sea in tropical and sub-tropical latitudes where they exist in conditions of high salinity, extreme tides, strong winds, high temperatures and muddy, anaerobic soils. There may be no other group of plants with such highly developed morphological and physiological adaptations to extreme conditions.

Because of their environment, mangroves are necessarily tolerant of high salt levels and have mechanisms to take up water despite strong osmotic potentials. Some also take up salts, but excrete them through specialized glands in the leaves. Others transfer salts into senescent leaves or store them in the bark or the wood. Still others simply become increasingly conservative in their water use as water salinity increases Morphological specializations include profuse lateral roots that anchor the trees in the loose sediments, exposed aerial roots for gas exchange and viviparous waterdispersed propagules.

Mangroves create unique ecological environments that host rich assemblages of species. The muddy or sandy sediments of the mangal are home to a variety of epibenthic, infaunal, and meiofaunal invertebrates Channels within the mangal support communities of phytoplankton, zooplankton and fish. The mangal may play a special role as nursery habitat for juveniles of fish whose adults occupy other habitats (e.g. coral reefs and seagrass beds).

Because they are surrounded by loose sediments, the submerged mangroves' roots, trunks and branches are islands of habitat that may attract rich epifaunal communities including bacteria, fungi, macroalgae and invertebrates. The aerial roots, trunks, leaves and branches host other groups of organisms. A number of crab species live among the roots, on the trunks or even forage in the canopy. Insects, reptiles, amphibians, birds and mammals thrive in the habitat and contribute to its unique character.

Living at the interface between land and sea, mangroves are well adapted to deal with natural stressors (e.g. temperature, salinity, anoxia, UV). However, because they live close to their tolerance limits, they may be particularly sensitive to disturbances like those created by human activities. Because of their proximity to population centers, mangals have historically been favored sites for sewage disposal. Industrial effluents have contributed to heavy metal contamination in the sediments. Oil from spills and from petroleum production has flowed into many mangals. These insults have had significant negative effects on the mangroves.

Habitat destruction through human encroachment has been the primary cause of mangrove loss. Diversion of freshwater for irrigation and land reclamation has destroyed extensive mangrove forests. In the past several decades, numerous tracts of mangrove have been converted for aquaculture, fundamentally altering the nature of the habitat. Measurements reveal alarming levels of mangrove destruction. Some estimates put global loss rates at one million ha y−1, with mangroves in some regions in danger of complete collapse. Heavy historical exploitation of mangroves has left many remaining habitats severely damaged.

These impacts are likely to continue, and worsen, as human populations expand further into the mangals. In regions where mangrove removal has produced significant environmental problems, efforts are underway to launch mangrove agroforestry and agriculture projects. Mangrove systems require intensive care to save threatened areas. So far, conservation and management efforts lag behind the destruction; there is still much to learn about proper management and sustainable harvesting of mangrove forests.

Mangroves have enormous ecological value. They protect and stabilize coastlines, enrich coastal waters, yield commercial forest products and support coastal fisheries. Mangrove forests are among the world's most productive ecosystems, producing organic carbon well in excess of the ecosystem requirements and contributing significantly to the global carbon cycle. Extracts from mangroves and mangrove-dependent species have proven activity against human, animal and plant pathogens. Mangroves may be further developed as sources of high-value commercial products and fishery resources and as sites for a burgeoning ecotourism industry. Their unique features also make them ideal sites for experimental studies of biodiversity and ecosystem function. Where degraded areas are being revegetated, continued monitoring and thorough assessment must be done to help understand the recovery process. This knowledge will help develop strategies to promote better rehabilitation of degraded mangrove habitats the world over and ensure that these unique ecosystems survive and flourish.

Biology of mangroves and mangrove Ecosystems

Changes in iron supply to oceanic plankton are thought to have a significant effect on concentrations of atmospheric carbon dioxide by altering rates of carbon sequestration, a theory known as the 'iron hypothesis' For this reason, it is important to understand the response of pelagic biota to increased iron supply Here we report the results of a mesoscale iron fertilization experiment in the polar Southern Ocean, where the potential to sequester iron-elevated algal carbon is probably greatest Increased iron supply led to elevated phytoplankton biomass and rates of photosynthesis in surface waters, causing a large drawdown of carbon dioxide and macronutrients, and elevated dimethyl sulphide levels after 13 days This drawdown was mostly due to the proliferation of diatom stocks But downward export of biogenic carbon was not increased Moreover, satellite observations of this massive bloom 30 days later, suggest that a sufficient proportion of the added iron was retained in surface waters Our findings demonstrate that iron supply controls phytoplankton growth and community composition during summer in these polar Southern Ocean waters, but the fate of algal carbon remains unknown and depends on the interplay between the processes controlling export, remineralisation and timescales of water mass subduction

A mesoscale phytoplankton bloom in the polar Southern Ocean stimulated by iron fertilization

A mesoscale phytoplankton bloom in the polar Southern Ocean stimulated by iron

Quaternary ammonium and tertiary sulfonium compounds in higher plants

The algal osmolyte dimethylsulphoniopropionate (DMSP) and its enzymatic cleavage product dimethylsulphide (DMS) contribute significantly to the global sulphur cycle, yet their physiological functions are uncertain. Here we report results that, together with those in the literature, show that DMSP and its breakdown products (DMS, acrylate, dimethylsulphoxide, and methane sulphinic acid) readily scavenge hydroxyl radicals and other reactive oxygen species, and thus may serve as an antioxidant system, regulated in part by enzymatic cleavage of DMSP. In support of this hypothesis, we found that oxidative stressors, solar ultraviolet radiation, CO(2) limitation, Fe limitation, high Cu(2+) (ref. 9) and H(2)O(2) substantially increased cellular DMSP and/or its lysis to DMS in marine algal cultures. Our results indicate direct links between such stressors and the dynamics of DMSP and DMS in marine phytoplankton, which probably influence the production of DMS and its release to the atmosphere. As oxidation of DMS to sulphuric acid in the atmosphere provides a major source of sulphate aerosols and cloud condensation nuclei, oxidative stressors--including solar radiation and Fe limitation--may be involved in complex ocean atmosphere feedback loops that influence global climate and hydrological cycles.

An antioxidant function for DMSP and DMS in marine algae

Marine plankton use a variety of defences against predators, some of which affect trophic structure and biogeochemistry1. We have previously shown2 that, during grazing by the protozoan Oxyrrhis marina on the alga Emiliania huxleyi, dimethylsulphoniopropionate (DMSP) from the prey is converted to dimethyl sulphide (DMS) when lysis of ingested prey cells initiates mixing of algal DMSP and the enzyme DMSP lyase. Such a mechanism is similar to macrophyte defence reactions3,4. Here we show that this reaction deters protozoan herbivores, presumably through the production of highly concentrated acrylate, which has antimicrobial activity5. Protozoan predators differ in their ability to ingest and survive on prey with high-activity DMSP lyase, but all grazers preferentially select strains with low enzyme activity when offered prey mixtures. This defence system involves investment in a chemical precursor, DMSP, which is not self-toxic and has other useful metabolic functions. We believe this is the first report of grazing-activated chemical defence in unicellular microorganisms.

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Grazing-activated chemical defence in a unicellular marine alga

Ecophysiology of polar algae

It has long been recognized that algae play a highly significant role in the global biogeochemical cycles of oxygen, carbon, nitrogen, phosphorus, and sulfw. The key chemical compounds driving these cycles are often low molecular weight and/or volatile species. In the case of sulfur, a dominant compound is dimethyl sulfide (DMS), which derives from dimethylsulfoniopropionate (DMSP) , an organic osmolyte that acts as a compatible solute in algal cells. High concentrations of DMS and DMSP can be found in areas of high algal productivity, such as marine phytoplankton blooms. In recent years research has unveiled greater detail concerning the biosynthesis and turnover of DMSP, DMS, and related compounds in the biosphere and with respect to the role of DMS in the global sulfur cycle and climate. In this minireview our aim is to concentrate mainly on these more recent discoveries.

Algal production of dimethyl sulfide and its atmospheric role1

SUMMARY The unicellular marine algae, Phaeodactylum tricornutum Bohlin, Cyclotella cryptica Reimann and Cyclotella meneghiniana Kutzing (Bacillariophyceae) and Porphyridium aerugineum Geitler (Rhodophyceae) synthesized and accumulated glycine betaine and proline in response to increases of the NaCl concentration (150 to 1000 mol m-3 NaCl) of the growth medium. C. cryptica and C. meneghiniana also synthesized and accumulated homarine (N-methyl picolinic acid betaine). Both P. tricornutum and P. aerugineum synthesized increasing amounts of intracellular glycerol and P. aerugineum also formed the heteroside, floridoside [O-a-Dgalactopyranosyl (1 -+ 2)-glycerol], in response to the elevated salinities. No major low molecular weight carbohydrates were found in Cyclotella. Sucrose was not detected in the algal extracts. Only P. tricornutum synthesized the tertiary sulphonium compound, ,-dimethylsulphoniopropionate (DMSP), and the quantity of this solute in the alga was dependent on the amount of NaCl in the medium. Intracellular K+ concentrations in the algae were three to six times greater than those of Na+. Increases of the salinity of the media led to the uptake and accumulation of K+ by the cells, and smaller increases of Na+ and Cl- and loss of intracellular NO3-. The inorganic cations Na+ and K+, with their accompanying anions, and the estimated organic solutes could largely account for the osmotic balance of P. tricornutum and P. aerugineum.

Osmotic adjustment in marine eukaryotic algae: the role of inorganic ions, quaternary ammonium, tertiary sulphonium and carbohydrate solutes

We characterised and compared dimethylsulfoniopropionate (DMSP) lyase isozymes in crude extracts of 6 axenic Emiliania huxleyi cultures (CCMP 370, 373, 374, 379, 1516, and strain L). This enzyme cleaves DMSP to form dimethyl sulfide (DMS), acrylate and a proton, but the function of this reaction in algae is still poorly understood. Most of the cultures produced high concentrations of intracellular DMSP, which was constant over the growth cycle and ranged from 157 to 242 mM, except for 1516 which had 50 mM DMSP cell-1. Extracts of all strains produced DMS from exogenous DMSP in vitro. DMSP lyases appeared constitutive, but enzyme activity and behaviour varied greatly among strains, and did not correlate with intracellular DMSP concentration. Strains 373 and 379 showed high DMSP lyase activities (12.5 and 6.1 fmol DMS cell-1 min-1, respectively), whereas DMS production was more than 100-fold lower in 370, 374, 1516 and L. This difference was intrinsic and the general pattern of high- and low-activity strains remained true over more than a 1 yr cultivation period. The cleavage reaction was optimal at pH 6 in the strains with high lyase activity and pH 5 was optimal for 374, 1516 and L. Strain 370 showed increasing activity with increasing pH. Experiments with additions of 0.125 to 2 M NaCl indicated halotolerant DMSP lyases in 373, 379 and 374. However, the halophilic DMSP lyases in 370 and L required 1 M NaCl addition for optimal DMSP cleavage, and 1516 showed optimal activity at 2 M NaCl. These results suggest that there are several structurally different DMSP lyase isozymes within E. huxleyi. However, it cannot be ruled out that varying concentrations of DMSP lyase per cell may have contributed to the differences in enzyme activity per cell. Comparison with other algal taxa indicates several families of DMSP lyases, hinting at possibly different cellular locations and functions, and varying DMS production under natural conditions.

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Gunter O. Kirst

Papers

Grazing-activated chemical defence in a unicellular marine alga

Ecophysiology of polar algae

Algal production of dimethyl sulfide and its atmospheric role1

Osmotic adjustment in marine eukaryotic algae: the role of inorganic ions, quaternary ammonium, tertiary sulphonium and carbohydrate solutes

Partial characterisation of dimethylsulfoniopropionate (DMSP) lyase isozymes in 6 strains of Emiliania huxleyi