Salt crystal deposition as a reversible mechanism to enhance photoprotection in black mangrove

Mangrove forests are ecosystems made up of several woody plants living in saline coastal sedimentary habitats. In order to deal with the high salinity of the substrate, mangrove trees possess a number of different mechanisms to exclude, sequestrate or excrete the excess of salt. The black mangrove (Avicennia germinans L.), one of the dominant species in Central America, is characterized by high levels of salt excretion through epidermal glands. In this study, our aim was to examine whether, apart from its obvious role in salt tolerance, the formation of salt crystals on the upper leaf surface of black mangrove might represent an unusual and dynamic photoprotection mechanism. For this purpose, the reflection of light and a number of physiological parameters were studied during the dry and rainy seasons in black mangroves growing in the Juan Venado Island Nature Reserve (Nicaragua). Excreted salt increased the reflectance of the leaf surface mainly in the blue and red regions of the spectrum. By removing salt crust from the leaf surface, we demonstrated that during the most stressful periods (dry season at noon), this feature allowed leaves to maintain a higher photochemical efficiency and a lower leaf temperature as compared to uncovered leaves. Furthermore, this mechanism is fully reversible when conditions become more favorable, as salt crystals dissolve, forming drops. Thus, while being a detoxification mechanism developed mainly to avoid osmotic imbalance in the tissues, the excretion of salt through the leaves in black mangroves is an example of “exaptation”, as it has positive collateral effects on the photosynthetic performance of the plant, protecting A. germinans from overheating and photodamage during the harsher periods.     

The Contribution of Mangrove Expansion to Salt Marsh Loss on the Texas Gulf Coast

Landscape-level shifts in plant species distribution and abundance can fundamentally change the ecology of an ecosystem. Such shifts are occurring within mangrove-marsh ecotones, where over the last few decades, relatively mild winters have led to mangrove expansion into areas previously occupied by salt marsh plants. On the Texas (USA) coast of the western Gulf of Mexico, most cases of mangrove expansion have been documented within specific bays or watersheds. Based on this body of relatively small-scale work and broader global patterns of mangrove expansion, we hypothesized that there has been a recent regional-level displacement of salt marshes by mangroves. We classified Landsat-5 Thematic Mapper images using artificial neural networks to quantify black mangrove (Avicennia germinans) expansion and salt marsh (Spartina alterniflora and other grass and forb species) loss over 20 years across the entire Texas coast. Between 1990 and 2010, mangrove area grew by 16.1 km², a 74% increase. Concurrently, salt marsh area decreased by 77.8 km², a 24% net loss. Only 6% of that loss was attributable to mangrove expansion; most salt marsh was lost due to conversion to tidal flats or water, likely a result of relative sea level rise. Our research confirmed that mangroves are expanding and, in some instances, displacing salt marshes at certain locations. However, this shift is not widespread when analyzed at a larger, regional level. Rather, local, relative sea level rise was indirectly implicated as another important driver causing regional-level salt marsh loss. Climate change is expected to accelerate both sea level rise and mangrove expansion; these mechanisms are likely to interact synergistically and contribute to salt marsh loss.     

Fungal diversity and plant disease in mangrove forests: salt excretion as a possible defense mechanism

Keywords. Salt excretion in leaves of some mangrove species may serve as an important defense against fungal attack, reducing the vulnerability of typically high-density, monospecific forest stands to severe disease pressure. In field surveys of a Caribbean mangrove forest in Panama, Avicennia germinans suffered much less damage from foliar diseases than did Laguncularia racemosa or Rhizophora mangle. Similarly, Avicennia leaves supported the least superficial fungal growth, endophytic colonization, and diversity, followed by Laguncularia and Rhizophora. Host specificity of leaf-colonizing fungi was greater than expected at random. We hypothesize that the different salt tolerance mechanisms in the three mangrove species may differentially regulate fungal colonization. The mangroves differ in their salt tolerance mechanisms such that Avicennia (which excretes salt through leaf glands) has the highest salinity of residual rain water on leaves, Laguncularia (which accumulates salt in the leaves) has the greatest bulk salt concentration, and Rhizophora (which excludes salt at the roots) has little salt associated with leaves. The high salt concentrations associated with leaves of Avicennia and Laguncularia, but not the low salinity of Rhizophora, were sufficient to inhibit the germination of many fungi associated with mangrove forests.     

Biotic interactions mediate the expansion of black mangrove (Avicennia germinans) into salt marshes under climate change

Many species are expanding their distributions to higher latitudes due to global warming. Understanding the mechanisms underlying these distribution shifts is critical for better understanding the impacts of climate changes. The climate envelope approach is widely used to model and predict species distribution shifts with changing climates. Biotic interactions between species, however, may also influence species distributions, and a better understanding of biotic interactions could improve predictions based solely on climate envelope models. Along the northern Gulf of Mexico coast, USA, subtropical black mangrove (Avicennia germinans) at the northern limit of its distribution grows sympatrically with temperate salt marsh plants in Florida, Louisiana, and Texas. In recent decades, freeze‐free winters have led to an expansion of black mangrove into salt marshes. We examined how biotic interactions between black mangrove and salt marsh vegetation along the Texas coast varied across (i) a latitudinal gradient (associated with a winter‐temperature gradient); (ii) the elevational gradient within each marsh (which creates different marsh habitats); and (iii) different life history stages of black mangroves (seedlings vs. juvenile trees). Each of these variables affected the strength or nature of biotic interactions between black mangrove and salt marsh vegetation: (i) Salt marsh vegetation facilitated black mangrove seedlings at their high‐latitude distribution limit, but inhibited black mangrove seedlings at lower latitudes; (ii) mangroves performed well at intermediate elevations, but grew and survived poorly in high‐ and low‐marsh habitats; and (iii) the effect of salt marsh vegetation on black mangroves switched from negative to neutral as black mangroves grew from seedlings into juvenile trees. These results indicate that the expansion of black mangroves is mediated by complex biotic interactions. A better understanding of the impacts of climate change on ecological communities requires incorporating context‐dependent biotic interactions into species range models.     

Phosphatidylglycerol Molecular Species in Chilling-Sensitive and Chilling-Resistant Populations of Avicennia germinans (L.) L

Phosphatidylglycerols (PG) from leaves and from roots of a chilling-sensitive(from Belize) and a chilling-tolerant (from Texas) populationof the black mangrove Avicennia germinans (L.) L were comparedwith respect to both their fatty acid and molecular speciescompositions. There were distinct differences between the twopopulations, and also between root and leaf PG of the same population.In leaf extracts the combined totals of palmitic acid and trans-3-hexadecenoicacid—two components thought to be correlated with chillingsensitivity—were not significantly different between thetwo populations. However, the PG in which only those fatty acidswere present, (i.e., dipalmitoyl and l-palmitoyl-2-(trans-3-hexadecenoyl)PG,accounted for a significantly greater proportion of the totalPG in leaves of the chilling-sensitive plants than in the moreresistant population. This trend in molecular species compositionwas not maintained in roots, where PGs were minor components.The findings are in only partial agreement with current ideasconcerning the possible role of specific lipid molecules ofchloroplast membranes in chilling tolerance. (Received May 28, 1984; Accepted September 3, 1984)     

Salt Tolerance in the Triticeae: K/Na Discrimination in Barley

Concentrations of ions were measured in the youngest fully-expandedleaves of Triticum aestivum, T. durum, Hordeum vulgare, H. spontaneum,Secale cereale, and Aegilops squarrosa accessions grown in hydroponicculture in the presence of salt (NaCl+CaCl₂). Triticum aestivum,Secale cereale, and Ae. squarrosa had the low leaf Na and highleaf K concentrations typical of plants which contain the enhancedK/Na discrimination character originally found in Ae. squarrosa.T. durum and the Hordeum species did not have this character.The better growth of H. vulgare than of T. durum with similarsalt concentrations in the youngest fully-expanded leaves maybe a result of better compartmentation of Na, Cl, and K betweendifferent tisssues or between different compartments withincells. The enhanced K/Na discrimination character was expressedin disomic addition lines of H. vulgare chromosomes in Triticumaestivum. The H. vulgare variety Herta and its slender mutantboth had similar leaf cation concentrations, although they differedin growth rate when grown at 60 mol m^–3 NaCl. H. vulgareand T. durum seedlings grown in the absence of monovalent cationsaccumulated more ²²Na in their shoots than seedlings of otherspecies when incubated in 1.0 mol m^–3 NaCl labelled with²²Na. Key words: Salt, ion transport, I genome, barley, wheat     

Away‐field advantage: mangrove seedlings grow best in litter from other mangrove species

Plant community composition can impact ecosystem processes via litter feedbacks. Species variation in litter quality may generate different patterns of nutrient supply for plants that are dependent on litter inputs. However, it is not known whether plants grow faster in their own litter, litter from other species, or in litter mixtures from multiple species. To test whether litter identity and mixture status influenced mangrove seedling growth, biomass allocation, and stoichiometry, we performed mesocosm experiments. Two species of mangrove seedlings, Avicennia germinans, black mangrove and Rhizophora mangle, red mangrove, were exposed to all possible combinations of three mangrove litter types and were isolated from all other nutrient inputs. Litter treatments significantly altered seedling growth. Seedlings from both mangrove species grew most rapidly in litter from a different species rather than their own, irrespective of litter chemical quality, decomposition rate, and nitrogen release. Litter mixtures from white and black mangroves caused black mangroves to grow 65% more than expected. Litter treatments did not impact seedling root:shoot ratios or tissue C:N. Our finding that seedlings grow best in litter from other species may indicate a mechanism that helps sustain the coexistence of dominant species.     

Herbivore feeding preferences as measured by leaf damage and stomatal ingestion: a mangrove crab example

The diet of the mangrove crab, Aratus pisonii, was assessed by determining the percent of damaged leaves at selected mangrove communities and by examining herbivore gut contents. This study compared the utility of both methods and tested if comparable levels of damage and dietary preference occurred using the methods. Percent of damaged leaves was determined for the three species of mangroves within Tampa Bay, FL, USA, including: the red, black, and white mangroves (Rhizophora mangle, Avicennia germinans, and Laguncularia racemosa, respectively) in four 5×10-m quadrats during summer 2001. For each species, in each of the quadrats, 200 leaves per tree were assessed for the presence or absence of crab damage. A. pisonii were sampled from the same quadrats from which leaf damage data were collected. Stomach contents were dissected and food items were classified into a number of categories.Species damaged and preferred were determined by comparing relative numbers of mangrove leaf stomata from the three mangrove species in gut contents. Results suggested that both methods provide similar estimates of preference. R. mangle leaves were preferred over those of A. germinans and L. racemosa. The percent of R. mangle leaves with damage was about 20-30 times greater than the other species, and R. mangle leaf stomata were 3 to 20 times more abundant in crab guts compared to leaf stomata of the other species. Gut contents indicated that A. pisonii is omnivorous, that average-sized adult crabs (1.4-1.7-cm width) prefer R. mangle, and that the diet of males is more varied than of females. While use of both percent leaf damage and crab gut contents reliably indicates preference, gut contents may describe better the actual diet and elucidate trends for different size or sex classes within a population.     

Epidermal Peels of Avicennia germinans (L.) Stearn: A Useful System to Study the Function of Salt Glands

Adaxial peels were made from fully expanded leaves of Avicenniagerminans (L). Stearn. The peels consisted of the epidermis,epidermal salt glands, and underlying hypodermal cells. Electronmicroscopic examinations showed that the structural integrity(except for the innermost hypodermal cells) of all cell typeswas not altered in making the peels. When the peels were floatedon salt solutions, the glands were shown to be functionallycompetent in secretion. Secretion also occurred when the peelswere floated on distilled water, presumably from salts storedin the hypodermis. Secretion occurred in the dark and sinceall cell types in the peels lacked chloroplasts, glandular functioncould not be directly coupled energetically to photosynthesis.However, secretion was shown to be temperature dependent andinhibited by azide and dinitrophenol, which indicates that theenergy cost underlying secretion is mitochondrial and most likelycoming from the mitochondria-enriched gland cells. Inhibitorsknown to affect membrane H⁺ ATPase activity and membrane transportalso inhibited secretion, indicating that membrane transportis probably the primary mechanism underlying secretion. Lanthanum,a membrane calcium antagonist, also inhibited secretion. Avicennia germinans (Avicenniaccac), salt glands, metabolic inhibitors, ultrastructure, secretion     

Mangrove litter dynamics in La Mancha Lagoon,Veracruz, Mexico

In the mangrove surrounding the coastal lagoon of La Mancha, Veracruz, Mexico, we studied litter fall, litter standing crop, and turnover rates in four different mangrove settings, based on the ecological classification of Lugo and Snedaker (1974). We studied those three prominent ecological processes at the basin, fringe and riverine mangrove settings, being the last one a relict riverine stand. The aim was to describe and compare litter dynamics among mangrove types in a lagoon with an ephemeral inlet, as a way of understanding functional heterogeneity within this coastal ecosystem. The daily average values of litter fall were different (P < 0.01) among mangrove site basin I, formed by Avicennia germinans and Rhizophora mangle (2.35 g/m²/day); basin II, formed by Laguncularia racemosa, Avicennia germinans, and Rhizophora mangle (2.93 g/m²/day); fringe with Rhizophora mangle (2.13 g/m²/day); and relic riverine, also with R. mangle (4.70 g/m²/day). The amount of litter standing crop was different among sites (P < 0.001), and also between the dry and rainy season, for each mangrove type (P < 0.001). Turnover ratios were higher in basin I and basin II sites (6.34 and 7.44 times per year) than in relic riverine and fringe mangroves (1.49 and 2.39 times per year). Interstitial salinity and sediment nutrients varied among mangrove types and could influence litter production. Since this lagoon has an ephemeral inlet, continuous inundation throughout 7–8 months per year has an important effect on litter dynamics.     

Salt Tolerance in the Triticeae: Leymus sabulosus

Elymus dahuhcus, Leymus giganteus, L. angustus, L. sabulosusand, to a lesser extent, L. triticoides, were found to tolerate200 mol m^–3 NaCl in solution culture. Elymus dahuricusdiffered from the Leymus species in its ion-uptake characteristics,showing a greater uptake of Cl and Na and a greater loss ofK from the shoots. In a more detailed experiment on Leymus sabulosusit was found that transpiration rates altered rapidly in responseto changes in external salinity whereas the accumulation ofNa and Cl in the leaves exhibited a lag of several days. Insalt stressed L. sabulosus Cl partially replaced the high levelsof nitrate found in the leaves of control plants. Glycinebetainelevels increased in the leaves from 8.0 mol m^–3 plantsap in the controls to 28 mol m^–3 plant sap at 250 molm^–3 NaCl. Key words: Salt stress, Transpiration, Solute accumulation, Leymus     

Is the central‐marginal hypothesis a general rule? Evidence from three distributions of an expanding mangrove species,Avicennia germinans (L.) L

The central‐marginal hypothesis (CMH) posits that range margins exhibit less genetic diversity and greater inter‐population genetic differentiation compared to range cores. CMH predictions are based on long‐held “abundant‐centre” assumptions of a decline in ecological conditions and abundances towards range margins. Although much empirical research has confirmed CMH, exceptions remain almost as common. We contend that mangroves provide a model system to test CMH that alleviates common confounding factors and may help clarify this lack of consensus. Here, we document changes in black mangrove (Avicennia germinans) population genetics with 12 nuclear microsatellite loci along three replicate coastlines in the United States (only two of three conform to underlying “abundant‐centre” assumptions). We then test an implicit prediction of CMH (reduced genetic diversity may constrain adaptation at range margins) by measuring functional traits of leaves associated with cold tolerance, the climatic factor that controls these mangrove distributional limits. CMH predictions were confirmed only along the coastlines that conform to “abundant‐centre” assumptions and, in contrast to theory, range margin A. germinans exhibited functional traits consistent with greater cold tolerance compared to range cores. These findings support previous accounts that CMH may not be a general rule across species and that reduced neutral genetic diversity at range margins may not be a constraint to shifts in functional trait variation along climatic gradients.     

The interplay between mangroves and saltmarshes at the transition between temperate and subtropical climate in Florida

The interplay between mangroves and saltmarshes at the temperate to subtropical transition in Florida results in dramatic changes to the appearance of the coastal landscape. In the 1980s, freezes killed entire mangrove forests dominated by black mangroves, Avicennia germinans (L.). Following the freezes, saltmarshes dominated by smooth cordgrass, Spartina alterniflora Loisel, revegetated the intertidal zone. After a decade of mild winters, however, mangroves are beginning to reclaim the area. The rate of mangrove expansion was determined by comparing aerial photography (change from 1995 to 1999), and from monitoring transects (over a 3 year period) on three of the Cedar Keys, Florida (Lat. 29°08′). The rate of mangrove expansion varied among islands, and the mechanism of expansion ranged from propagule-trapping by saltmarshes along the edges of mangrove clumps to widespread dispersal and growth of existing or newly imported propagules. A freeze occurred during the study, which may have set back mangrove expansion by defoliating mangrove trees and resetting mangrove reproduction. Mangrove expansion was projected to take 20–30 years for complete seedling cover. Given the possibility of global climate change and its potential influence on the distribution of coastal vegetation, the timeframes and implications to coastal wetland ecosystems involved in this regular interplay will provide valuable baseline information for future studies.     

Where temperate meets tropical: multi-factorial effects of elevated CO2, nitrogen enrichment, and competition on a mangrove-salt marsh community

Our understanding of how elevated CO₂ and interactions with other factors will affect coastal plant communities is limited. Such information is particularly needed for transitional communities where major vegetation types converge. Tropical mangroves (Avicennia germinans) intergrade with temperate salt marshes (Spartina alterniflora) in the northern Gulf of Mexico, and this transitional community represents an important experimental system to test hypotheses about global change impacts on critical ecosystems. We examined the responses of A. germinans (C₃) and S. alterniflora (C₄), grown in monoculture and mixture in mesocosms for 18 months, to interactive effects of atmospheric CO₂ and pore water nitrogen (N) concentrations typical of these marshes. A. germinans, grown without competition from S. alterniflora, increased final biomass (35%) under elevated CO₂ treatment and higher N availability. Growth of A. germinans was severely curtailed, however, when grown in mixture with S. alterniflora, and enrichment with CO₂ and N could not reverse this growth suppression. A field experiment using mangrove seedlings produced by CO₂‐ and N‐enriched trees confirmed that competition from S. alterniflora suppressed growth under natural conditions and further showed that herbivory greatly reduced survival of all seedlings. Thus, mangroves will not supplant marsh vegetation due to elevated CO₂ alone, but instead will require changes in climate, environmental stress, or disturbance to alter the competitive balance between these species. However, where competition and herbivory are low, elevated CO₂ may accelerate mangrove transition from the seedling to sapling stage and also increase above‐ and belowground production of existing mangrove stands, particularly in combination with higher soil N.     

Monitoring of black mangrove restoration with nursery-reared seedlings on an arid coastal lagoon

Black mangrove (Avicennia germinans) seedlings (n=555) were grown from field-collected propagules for 3 months in a new type of terrestrial nursery. They were grown in clusters of five plants, and then they were transplanted to a clear-cut zone in a lagoon fringed by a mangrove forest at Laguna de Balandra, Baja California Sur, Mexico. Survival and plant development of transplants were monitored at 6-monthly intervals for 2 years. After 1 month, the survival of seedlings was 96%, later stabilizing at approximately 77%. After 24 months, 74% of the plants were still alive. The best cluster, showing maximum growth under mangrove swamp conditions in this arid zone, was a two-plant cluster. The lagoon has a low natural regeneration rate of 48 plants per 350 m² per 6 years of monitoring. This study shows the feasibility of restoring destroyed arid-coast lagoons with black mangroves.     

First report of Avicennia germinans (Acanthaceae) as a larval host of Junonia litoralis (Nymphalidae) causing higher foliar damage in seedlings than trees

Herbivory is considered an important biotic interaction in mangroves. Nevertheless, detailed information on specific plant–herbivore interactions that might have considerable influence in ecological and evolutionary processes is still very poor and fragmented. Herbivory damage was quantified during December 2015 in seedlings and trees in monospecific stands of Avicennia germinans in Laguna Madre of Tamaulipas, Mexico. The use of this mangrove species as a larval host is well documented; however, this is the first report for Junonia litoralis. These lepidopteran larvae consumed, on average, 10.2% of the leaf area; but herbivory was higher on seedlings (mean 15%) than on trees (mean 2.3%). These values are comparable to estimations of 10% mean herbivory damage in other field studies. This indicates that herbivores cause greater damage in seedlings than in trees. Herbivore activity by J. litoralis only occurred in A. germinans and did not affect associated herbs or shrubs of other species. The close interaction between J. litoralis and A. germinans may have profound implications for ecological and evolutionary processes of mangroves and enables a better understanding of ecosystem function and its conservation. Further studies are needed to investigate such interactions and their implications including long-term monitoring of interstitial salinity and leaf chemistry at different stages of growth and maturity of the host species.     

The role of saltwort (Batis maritima L.) in regeneration of degraded mangrove forests

While saltwort (Batis maritima L.) is common in the fringe mangrove forests of southwest Florida, its role in regeneration of degraded mangrove communities is not known. Given the potential encroachment and subsequent degradation of mangrove communities by sea-level rise, it is important to quantify the effect of early-colonizing vegetation to early mangrove seedling survival. A greater number of mangrove seedlings were observed in existing B. maritima patches compared to surrounding mudflats. A planting experiment was designed to determine whether B. maritima was responsible for the observed pattern. Black mangrove (Avicennia germinans L.) seedlings, raised in a nursery, were planted in previously established B. maritima patches and on mudflats with and without nursery-raised B. maritima. There was significantly lower mortality of A. germinans seedlings when planted in existing B. maritima patches (69%), compared to seedlings planted on the mudflats (93%), demonstrating that existing B. maritima improved A. germinans seedling survival. Nursery-raised B. maritima had lower mortality on open mudflats (28%), suggesting that it can tolerate conditions, which make it an early colonizer of newly available habitats. The primary mechanism proposed for improving seedling success is a slight increase in elevation provided by the dense root network of established B. maritima. These findings have implications for scientists and managers anticipating the response of mangroves to sea-level rise.     

Differentiation between true mangroves and mangrove associates based on leaf traits and salt contents

Aims Mangrove species are classified as true mangroves and mangrove associates. However, as for some fringe species found mainly on the landward transitional zones of mangroves, no consensus among scientists could be reached in favor of this classification and much debate arises. We hypothesized that true mangroves differ from mangrove associates physiologically and ecologically in their ability to survive in mangrove environment.Methods To test this hypothesis, leaf structural traits and osmotic properties were used to describe variation in 33 mangrove species (17 true mangroves, 6 mangrove associates and 10 controversial species).Important findings Specific leaf area (SLA) of true mangroves as well as leaf nitrogen concentration on a leaf mass (Nmass) were lower than that of mangrove associates; leaf succulence was, in general, twice as high in true mangroves compared to mangrove associates; true mangroves accumulated 8–9 times more Na and Cl than mangrove associates and the former had K/Na ratios <0.5, but the latter had K/Na ratios>0.5. These results indicated that true mangroves differed reliably from mangrove associates in leaf traits and osmotic properties. True mangroves are true halophytes and mangrove associates are glycophytes with certain salt tolerance. Combining distribution pattern information, the 10 controversial species were reclassified.     

Will fluctuations in salt marsh–mangrove dominance alter vulnerability of a subtropical wetland to sea‐level rise?

To avoid submergence during sea‐level rise, coastal wetlands build soil surfaces vertically through accumulation of inorganic sediment and organic matter. At climatic boundaries where mangroves are expanding and replacing salt marsh, wetland capacity to respond to sea‐level rise may change. To compare how well mangroves and salt marshes accommodate sea‐level rise, we conducted a manipulative field experiment in a subtropical plant community in the subsiding Mississippi River Delta. Experimental plots were established in spatially equivalent positions along creek banks in monospecific stands of Spartina alterniflora (smooth cordgrass) or Avicennia germinans (black mangrove) and in mixed stands containing both species. To examine the effect of disturbance on elevation dynamics, vegetation in half of the plots was subjected to freezing (mangrove) or wrack burial (salt marsh), which caused shoot mortality. Vertical soil development was monitored for 6 years with the surface elevation table‐marker horizon system. Comparison of land movement with relative sea‐level rise showed that this plant community was experiencing an elevation deficit (i.e., sea level was rising faster than the wetland was building vertically) and was relying on elevation capital (i.e., relative position in the tidal frame) to survive. Although Avicennia plots had more elevation capital, suggesting longer survival, than Spartina or mixed plots, vegetation type had no effect on rates of accretion, vertical movement in root and sub‐root zones, or net elevation change. Thus, these salt marsh and mangrove assemblages were accreting sediment and building vertically at equivalent rates. Small‐scale disturbance of the plant canopy also had no effect on elevation trajectories—contrary to work in peat‐forming wetlands showing elevation responses to changes in plant productivity. The findings indicate that in this deltaic setting with strong physical influences controlling elevation (sediment accretion, subsidence), mangrove replacement of salt marsh, with or without disturbance, will not necessarily alter vulnerability to sea‐level rise.     

Changes in intracellular electrolyte concentrations during apoptosis induced by UV irradiation of human myeloblastic cells

Decreases in the intracellular concentrations of both K⁺ and Cl^– have been implicated in playing a major role in the progression of apoptosis, but little is known about the temporal relationship between decreases in electrolyte concentration and the key events in apoptosis, and there is no information about how such decreases affect different intracellular compartments. Electron probe X-ray microanalysis was used to determine changes in element concentrations (Na, P, Cl, and K) in nucleus, cytoplasm, and mitochondria in U937 cells undergoing UV-induced apoptosis. In all compartments, the initial stages of apoptosis were characterized by decreases in [K] and [Cl]. The largest decreases in these elements were in the mitochondria and occurred before the release of cytochrome c. Initial decreases in [K] and [Cl] also preceded apoptotic changes in the nucleus. In the later stages of apoptosis, the [K] continued to decrease, whereas that of Cl began to increase toward control levels and was accompanied by an increase in [Na]. In the nucleus, these increases coincided with poly(ADP-ribose) polymerase cleavage, chromatin condensation, and DNA laddering. The cytoplasm was the compartment least affected and the pattern of change of Cl was similar to those in other compartments, but the decrease in [K] was not significant until after active caspase-3 was detected. Our results support the concept that normotonic cell shrinkage occurs early in apoptosis, and demonstrate that changes in the intracellular concentrations of K and Cl precede apoptotic changes in the cell compartments studied. sodium; potassium; chloride; cell shrinkage