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

Recent progress in improving the salt tolerance of cultivated plants has been slow. Physiologists have been unable to define single genes or even specific metabolic processes that molecular biologists could target, or pinpoint the part of the plant in which such genes for salt tolerance might be expressed. While the physiological might be expressed. While the physiological processes are undoubtedly complex, faster progress on unraveling mechanisms of salt tolerance might be made if there were more effort to test hypotheses rather than to accumulate data, and to integrate cellular and whole plant responses. This article argues that salts taken up by the plant do not directly control plant growth by affecting turgor, photosynthesis or the activity of any one enzyme. Rather, the build-up of salt in old leaves hasten their death, and the loss of these leaves affects the supply of assimilates or hormones to the growing regions and thereby affects growth.

Physiological processes limiting plant growth in saline soils: some dogmas and hypotheses

Plants and their arbuscular mycorrhizal fungal symbionts interact in complex underground networks involving multiple partners. This increases the potential for exploitation and defection by individuals, raising the question of how partners maintain a fair, two-way transfer of resources. We manipulated cooperation in plants and fungal partners to show that plants can detect, discriminate, and reward the best fungal partners with more carbohydrates. In turn, their fungal partners enforce cooperation by increasing nutrient transfer only to those roots providing more carbohydrates. On the basis of these observations we conclude that, unlike many other mutualisms, the symbiont cannot be "enslaved." Rather, the mutualism is evolutionarily stable because control is bidirectional, and partners offering the best rate of exchange are rewarded.

https://science.sciencemag.org/content/sci/333/6044/880.full.pdf

Reciprocal Rewards Stabilize Cooperation in the Mycorrhizal Symbiosis

Summary Here, the coevolution of mycorrhizal fungi and roots is assessed in the light of evidence now available, from palaeobotanical and morphological studies and the analysis of DNA-based phylogenies. The first bryophyte-like land plants, in the early Devonian (400 million years ago), had endophytic associations resembling vesicular‐ arbuscular mycorrhizas (VAM) even before roots evolved. Mycorrhizal evolution would have progressed from endophytic hyphae towards balanced associations where partners were interdependent due to the exchange of limiting energy and nutrient resources. Most mycorrhizas are mutualistic, but in some cases the trend for increasing plant control of fungi culminates in the exploitative mycorrhizas of achlorophyllous, mycoheterotrophic plants. Ectomycorrhizal, ericoid and orchid mycorrhizas, as well as nonmycorrhizal roots, evolved during the period of rapid angiosperm radiation in the Cretaceous. It is hypothesised that roots gradually evolved from rhizomes to provide more suitable habitats for mycorrhizal fungi and provide plants with complex branching and leaves with water and nutrients. Selection pressures have caused the morphological divergence of roots with different types of mycorrizas. Root cortex thickness and exodermis suberization are greatest in obligately mycorrhizal plants, while nonmycorrhizal plants tend to have fine roots, with more roots hairs and relatively advanced chemical defences. Major coevolutionary trends and the relative success of plants with different root types are discussed.

Coevolution of roots and mycorrhizas of land plants

A comprehensive appraisal of the mycorrhizal literature provides data for 336 plant families representing 99% of flowering plants, with regard to mycorrhizas and other nutritional adaptations. In total, arbuscular (AM), orchid, ectomycorrhizas (EM) and ericoid mycorrhizas and nonmycorrhizal (NM) roots occur in 74%, 9%, 2%, 1% and 6% of Angiosperm species respectively. Many families of NM plants have alternative nutritional strategies such as parasitism, carnivory, or cluster roots. The remaining angiosperms (8%) belong to families reported to have both AM and NM species. These are designated as NM-AM families here and tend to occur in habitats considered non-conducive to mycorrhizal fungi, such as epiphytic, aquatic, extremely cold, dry, disturbed, or saline habitats. Estimated numbers of species in each category of mycorrhizas is presented with lists of NM and EM families. Evolutionary trends are also summarised by providing data on all clades and orders of flowering and non-flowering vascular plants on a global scale. A case study of Western Australian plants revealed that plants with specialised nutritional modes such as carnivory, cluster roots, or EM were much more diverse in this ancient landscape with infertile soils than elsewhere. Detailed information on the mycorrhizal diversity of plants presented here is linked to a website (mycorrhizas.info) to allow data to remain current. Over a century of research effort has resulted in data on mycorrhizal associations of >10,000 plant species that are of great value, but also somewhat of a liability due to conflicting information about some families and genera. It is likely that these conflicts result in part from misdiagnosis of mycorrhizal associations resulting from a lack of standardisation in criteria used to define them. Families that contain both NM and AM species provide a second major source of inconsistency, but even when these are excluded there is a ∼10% apparent error rate in published lists of mycorrhizal plants. Arbuscules are linked to AM misdiagnosis since they are used less often than vesicles to recognise AM associations in roots and apparently occur sporadically in NM plants. Key issues with the diagnosis of mycorrhizal plants are discussed using the Cyperaceae as a case study. Detailed protocols designed to consistently distinguish AM from endophytic Glomeromycotan Fungus Colonisation (GFC) are provided. This review aims to stimulate debate and provide advice to researchers delving into root biology.

Mycorrhizal associations and other means of nutrition of vascular plants: understanding the global diversity of host plants by resolving conflicting information and developing reliable means of diagnosis

Metal localization within and around roots of water hyacinth (Eichhornia crassipes) growing in a wetland receiving urban run-off was studied by energy dispersive X-ray microanalysis of sections from freeze-substituted roots. Sampling randomly from an order of magnitude gradient in metal concentrations (Cu and Pb) allowed us the opportunity to identify general patterns of metal localization. Iron was present at high levels at the root surface, and this may have been a root plaque as described for wetland plants with roots anchored in flooded soils. Iron levels decreased centripetally across the root and were higher in cell walls than within cells. Trace metals (Cu, Zn and Pb) were not localized at the root surface. In contrast with iron, trace metal levels increased centripetally across the root, tended to be higher inside cells and were highest within cells in the stele. Variability of localization was high for all metals analysed. Multivariate statistical analyses (principal components analysis and multidimensional scaling) were useful for identifying overall patterns in elemental distribution.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-3040.1999.00388.x

Metal localization in water hyacinth roots from an urban wetland

The influence of external phosphorus (P) on carbon (C) allocation and metabolism as well as processes related to P metabolism was studied in monoxenic arbuscular mycorrhiza cultures of carrot ( Daucus carota ). Fungal hyphae of Glomus intraradices proliferated from the solid minimal medium containing the colonized roots into C-free liquid minimal medium with different P treatments. The fungus formed around three times higher biomass in P-free liquid medium than in medium with 2.5 mminorganic P (high-P). Mycelium in the second experiment was harvested at an earlier growth stage to study metabolic processes when the mycelium was actively growing. P treatment influenced the root P content and [ 13 C]glucose administered to the roots 7 d before harvest gave a negative correlation between root P content and 13 C enrichment in arbuscular mycorrhiza fungal storage lipids in the extraradical hyphae. Eighteen percent of the enriched 13 C in extraradical hyphae was recovered in the fatty acid 16:1ω5 from neutral lipids. Polyphosphate accumulated in hyphae even in P-free medium. No influence of P treatment on fungal acid phosphatase activity was observed, whereas the proportion of alkaline-phosphatase-active hyphae was highest in high-P medium. We demonstrated the presence of a motile tubular vacuolar system in G. intraradices . This system was rarely seen in hyphae subjected to the highest P treatment. We concluded that the direct responses of the extraradical hyphae to the P concentration in the medium are limited. The effects found in hyphae seemed instead to be related to increased availability of P to the host root.

Phosphorus Effects on Metabolic Processes in Monoxenic Arbuscular Mycorrhiza Cultures

Leaf and other litter falling in a mangrove woodland was estimated by litter trapping for 13 months. Mean annual litter fall was 5.8 t ha-1 year-1 (dry matter) of which 79% was leaf material. Largest fortnightly collections of litter were found in the summer months, and little litter fell in winter. Data suggested that mangrove litter is rapidly incorporated into the rest of the estuarine ecosystem: the leaf litter layer in the swamp was estimated on two occasions (about 9 g m-2 in winter and about 62 g m-2 in summer, dry leaf material) and a litter-bag experiment gave a leaf decomposition half- time of about 8 weeks.

Litter Fall and Decompostion in a Mangrove Stand, Avicennia marina (Forsk.) Vierh., in Middle Harbour, Sydney

summary
In intact seedlings of Eucalyptus pilularis Stn. ectomycorrhizal with Pisolithus tinctorius (Pers.) Coker & Couch in growth pouches, and labelled photosynthetically with 14C4 more radioactivity was translocated into mycorrhizal than to non-mycorrhizal roots. Results from autoradiography of intact root systems were supported by data from scintillation counting of individual root tips. Greater 14C accumulation occurred in young than in older mycorrhizas. It is proposed that individual mycorrhizal roots act most strongly as carbon sinks early in their life and that there is a progressive reduction in the amount of photosynthate translocated to them as they age.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1989.tb00343.x

Distribution of photosynthetically fixed carbon within root systems of Eucalyptus pilularis plants ectomycorrhizal with Pisolithus tinctorius.

A method is described by which enzyme activity can be retained in plant and animal tissues embedded in glycol methacrylate at low temperature.

W. G. Allaway

Papers

Metal localization in water hyacinth roots from an urban wetland

Phosphorus Effects on Metabolic Processes in Monoxenic Arbuscular Mycorrhiza Cultures

Litter Fall and Decompostion in a Mangrove Stand, Avicennia marina (Forsk.) Vierh., in Middle Harbour, Sydney

Distribution of photosynthetically fixed carbon within root systems of Eucalyptus pilularis plants ectomycorrhizal with Pisolithus tinctorius.

Low temperature embedding in glycol methacrylate for enzyme histochemistry in plant and animal tissues.