Role of root hydrophobic barriers in salt exclusion of a mangrove plant Avicennia officinalis

Salt exclusion at the roots and salt secretion in the leaves were examined in a mangrove, Avicennia officinalis. The non‐secretor mangrove Bruguiera cylindrica was used for comparative study of hydrophobic barrier formation in the roots. Bypass flow was reduced when seedlings were previously treated with high salt concentration. A biseriate exodermis was detected in the salt‐treated roots, along with an enhanced deposition of hydrophobic barriers in the endodermis. These barriers reduced Na⁺ loading into the xylem, accounting for a 90–95% salt exclusion in A. officinalis. Prominent barriers were found in the roots of B. cylindrica even in the absence of salt treatment. A cytochrome P450 gene that may regulate suberin biosynthesis was up‐regulated within hours of salt treatment in A. officinalis roots and leaves, corresponding with increased suberin deposition. X‐ray microanalysis showed preferential deposition of Na⁺ and Cl^? in the root cortex compared with the stele, suggesting that the endodermis is the primary site of salt exclusion. Enhanced salt secretion and increased suberin deposition surrounding the salt glands were seen in the leaves with salt treatment. Overall, these data show that the deposition of apoplastic barriers increases resistance to bypass flow leading to efficient salt exclusion at the roots in mangroves.     

三种泌盐红树植物对盐胁迫的耐受性比较

在盐度 0、5、15、2 5和 35 (% )下种植泌盐红树植物老鼠 (Acanthus ilicifolius)、桐花树 (Aegiceras corniculatum)和白骨壤(Avicennia marina)的繁殖体 ,以繁殖体萌发、幼苗生长、叶片泌盐量、叶片组织液盐含量和蒸腾蒸发量为指标 ,比较其对盐胁迫的耐受性。盐度提高对胎生种类桐花树和白骨壤的萌根速率无显著影响 ,但高盐度明显抑制非胎生种类老鼠的萌根。白骨壤的萌苗率不受盐度影响 ,但 2 5以上的盐度导致桐花树和老鼠的萌苗率下降。在盐度范围 5～ 35内 ,白骨壤幼苗的茎高生长随盐度的增加而减少 ,但减少量比桐花树小 ,而老鼠的减少量最大。老鼠因盐度提高而导致的叶片长度的减少量最大。在盐度提高的情况下 3种植物均具有泌盐量增加的效应 ,在任一盐度下泌盐能力的顺序均为白骨壤 >桐花树 >老鼠。淡水培养时 ,3种红树植物的叶片组织液盐含量 (约 2 % )均高于环境盐度 0。在盐度范围 5～ 35内 ,白骨壤的叶片组织液盐含量维持在较稳定的水平 (4 .3%～ 5 .0 % ) ,桐花树的变化范围为 2 .4 %～ 4 .5 % ,老鼠 2 .3%～ 5 .3%。淡水培养时 ,3种植物的蒸腾蒸发量类似 ,但盐性条件下白骨壤的蒸腾蒸发量显著高于桐花树和老鼠。随着盐度的增加 ,老鼠的蒸腾蒸发量下降最多。这些结果均表     

Growth responses of the mangrove Avicennia marina to salinity: development and function of shoot hydraulic systems require saline conditions

Background and Aims Halophytic eudicots are characterized by enhanced growth under saline conditions. This study combines physiological and anatomical analyses to identify processes underlying growth responses of the mangrove Avicennia marina to salinities ranging from fresh- to seawater conditions.Methods Following pre-exhaustion of cotyledonary reserves under optimal conditions (i.e. 50 % seawater), seedlings of A. marina were grown hydroponically in dilutions of seawater amended with nutrients. Whole-plant growth characteristics were analysed in relation to dry mass accumulation and its allocation to different plant parts. Gas exchange characteristics and stable carbon isotopic composition of leaves were measured to evaluate water use in relation to carbon gain. Stem and leaf hydraulic anatomy were measured in relation to plant water use and growth.Key Results Avicennia marina seedlings failed to grow in 0–5 % seawater, whereas maximal growth occurred in 50–75 % seawater. Relative growth rates were affected by changes in leaf area ratio (LAR) and net assimilation rate (NAR) along the salinity gradient, with NAR generally being more important. Gas exchange characteristics followed the same trends as plant growth, with assimilation rates and stomatal conductance being greatest in leaves grown in 50–75 % seawater. However, water use efficiency was maintained nearly constant across all salinities, consistent with carbon isotopic signatures. Anatomical studies revealed variation in rates of development and composition of hydraulic tissues that were consistent with salinity-dependent patterns in water use and growth, including a structural explanation for low stomatal conductance and growth under low salinity.Conclusions The results identified stem and leaf transport systems as central to understanding the integrated growth responses to variation in salinity from fresh- to seawater conditions. Avicennia marina was revealed as an obligate halophyte, requiring saline conditions for development of the transport systems needed to sustain water use and carbon gain.     

Isolation and characterization of 10 new compound microsatellite markers for a mangrove tree species,Avicennia marina (Forsk.) Vierh. (Avicenniaceae)

Avicennia marina is an ecologically important mangrove tree species. We isolated 10 polymorphic microsatellite loci from this species using an improved technique. Our isolated loci provided compound microsatellite markers with polymorphism of two to six alleles per locus. The observed and expected heterozygosities ranged from 0.025 to 0.625 and from 0.096 to 0.767, respectively. These markers would be the useful tools for researching on the genetic diversity and population genetic structure of A. marina.     

Growth and respiration in two mangrove species at a range of salinities

Growth and dark respiration rates were measured in leaves and roots of seedlings of Avicennia marina (Forsk.) Vierh, (grey mangrove), and Aegiceras corniculatum (L.) Blanco (river mangrove). Plants were grown in a soil mixture at ambient temperatures and watered with 0.25 and 100% sea-water. Oxygen uptake was measured in excised root and leaf samples. In both species growth was maximal in 25% sea-water, and root respiration was lowest in 100% sea-water. Differences were found between the two species in the responses of leaf respiration to salinity. In A. corniculatum leaf respiration was raised in both 25 and 100% sea-water, while in A. marina only leaves in 100% sea-water showed higher rates of respiration. These results are consistent with the view that A. marina is the more salt-tolerant of the two species. In A. corniculatum the respiration rates of the hypocotyl were also measured, and were much higher in 100% sea-water than in the other two treatments. The results suggest that at high salinities there is a high metabolic cost in the shoots of both species, and that at such salinities rates of root respiration may be limited by the supply of substrate from the shoots.     

Microsatellite analysis of genetic structure in the mangrove species Avicennia marina (Forsk.) Vierh. (Avicenniaceae)

The level of genetic variation throughout the entire worldwide range of the mangrove species Avicennia marina (Forsk.) Vierh. was examined using microsatellite markers. Three microsatellite loci detected high levels of allelic diversity (70 alleles in total), essential for an accurate estimation of population genetic parameters. The informativeness of the microsatellite loci tended to increase with increasing average number of repeats. The levels of heterozygosity detected for each population, over all loci, ranged from 0.0 to 0.8, with an average of 0.407, indicating that some populations had little or no genetic variation, whereas others had a large amount. Populations at the extremes of the distribution range showed reduced levels of heterozygosity, and significant levels of inbreeding. This is not unexpected as these populations may be subject to founder effects and environmental constraints. The presence of genetic structure was tested in A. marina populations using three models: (i) a single panmictic model; (ii) the discrete subpopulation model; and (iii) the isolation by distance model. The discrete subpopulations model was supported by the overall measures of population differentiation based on the infinite alleles model (F-statistics), and the stepwise mutation model (R statistics). In addition, an analysis of molecular variance (AMOVA), using both theoretical models, found that most of the variation was between populations (41-71%), and within individuals in the total population (31-49%). There was little variation among individuals within populations (0-10%). There was no significant isolation by distance. The high levels of genetic differentiation observed among populations of A. marina may be due to environmental and ecological factors, particularly past sea level and climatic changes.     

Effect of salt stress on the metabolism of ethanolamine and choline in leaves of the betaine-producing mangrove species Avicennia marina

Glycinebetaine synthesis from [methyl-14C]choline and [1,2-14C]ethanolamine in leaf disks of Avicennia marina, was increased by salt stress (250 and 500 mM NaCl). After 18 h incubation with [methyl-14C]choline, phosphocholine and CO(2) were found to be heavily labelled. Phosphocholine contained 39% of the total radioactivity taken up by non-salinised (control) leaf disks and 15% of the total for salinised leaf disks stressed with 500 mM NaCl. Eighteen and 49% of the radioactivity absorbed by control and salinised disks, respectively, were released as CO(2). Metabolic studies of [1,2-14C]ethanolamine revealed that the radioactivity taken up by the leaf disks was recovered as the following compounds after 18 h: phosphorylated compounds (mainly phosphoethanolamine, phosphodimethylethanolamine and phosphocholine) (40-50%); choline (1-2%); glycinebetaine (3-5%); lipids (20-28%); CO(2) (6-10%). Unlike glycinebetaine, incorporation into phosphorylated compounds and lipids were reduced by salt stress. Incorporation of [methyl-14C]S-adenosyl-L-methionine (SAM) into choline, phosphocholine and glycinebetaine in leaf disks was stimulated by salt stress. In vitro activities of adenosine kinase and adenosine nucleosidase, which are implicated in stimulating the SAM regeneration cycle, increased after the leaf disks were incubated with 250 and 500 mM NaCl for 18 h. Changes in metabolism involving choline and glycinebetaine due to salt stress are discussed.     

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.     

Ultrastructural changes accompanying secretion and cell death in the molting glands of an insect (Oncopeltus)

During the fifth (last) larval instar of Oncopeltus fasciatus, morphological changes in the molting glands associated with ecdysone secretion include an increase in cytoplasmic volume relative to that of the nucleus, increased amounts of rough endoplasmic reticulum and mitochondria, and the formation of deep infoldings of the plasma membrane. On the sixth day of the fifth instar large electron-lucent areas become apparent beneath the basement membrane; however, the glands remain intact until the seventh (last) day of the instar when a dramatic fragmentation of the cytoplasm, and condensation and fragmentation of the nucleus are observed. It is likely that such changes occur rapidly, just prior to the time of ecdysis to an adult. Cell death in the molting glands of Oncopeltus is markedly different from that described for the molting glands of other insect species in that autophagic vacuoles are not observed prior to a complete loss of cellular integrity.     

Winter climate change and coastal wetland foundation species: salt marshes vs. mangrove forests in the southeastern United States

We live in an era of unprecedented ecological change in which ecologists and natural resource managers are increasingly challenged to anticipate and prepare for the ecological effects of future global change. In this study, we investigated the potential effect of winter climate change upon salt marsh and mangrove forest foundation species in the southeastern United States. Our research addresses the following three questions: (1) What is the relationship between winter climate and the presence and abundance of mangrove forests relative to salt marshes; (2) How vulnerable are salt marshes to winter climate change‐induced mangrove forest range expansion; and (3) What is the potential future distribution and relative abundance of mangrove forests under alternative winter climate change scenarios? We developed simple winter climate‐based models to predict mangrove forest distribution and relative abundance using observed winter temperature data (1970–2000) and mangrove forest and salt marsh habitat data. Our results identify winter climate thresholds for salt marsh–mangrove forest interactions and highlight coastal areas in the southeastern United States (e.g., Texas, Louisiana, and parts of Florida) where relatively small changes in the intensity and frequency of extreme winter events could cause relatively dramatic landscape‐scale ecosystem structural and functional change in the form of poleward mangrove forest migration and salt marsh displacement. The ecological implications of these marsh‐to‐mangrove forest conversions are poorly understood, but would likely include changes for associated fish and wildlife populations and for the supply of some ecosystem goods and services.     

Fine structural aspects of silk secretion in a spider. II. Conduction in the pyriform glands

The excretory duct of pyriform glands in Araneus diadematus is connected to the secretory sac through an intermediary cell ring. Apices of these cells bear thick, long microvilli and cytoplasmic extensions containing microtubules in bundles, some of which are derived from normal basal bodies. These finger-like extensions lie between the cuticular intima and the secretory product; they are thought to protect the intima and to initiate moulding of the silk thread. Structural features of the duct cells suggest that the latter play a role in the control of the water content of the silk glue which is restricted to the last portion of the duct where numerous nerve endings are inserted between cells. It is evident that duct structure and chemical and physical characteristics of silk are correlated in all spider silk glands.     

Comparison of Distichlis spicata and Suaeda aegyptiaca in response to water salinity: Candidate halophytic species for saline soils remediation

Distichlis spicata and Suaeda aegyptiaca are two potential halophytic plant species for bioremediation of salt degraded soils, and development of saline agriculture. The physiological responses of the species to different levels of salinity (EC 0, 12, 24, 36, and 48 dS/m) in a controlled environment experiment were studied. Both species showed a high level of tolerance to elevated concentrations of salt in the irrigation water. The shoot fresh and dry weights in S. aegyptiaca increased till 36 dS/m and were sustained under 48 dS/m while in D. spicata, both parameters decreased as salinity increased. Glycine betaine accumulation did not change in D. spicata with increasing salinity, whereas proline content revealed a marked increase of 7.13 fold in 48 dS/m salinity compared to the control, which showed its critical osmoprotection role in the plant. In S. aegyptiaca, both osmolytes content significantly increased at high salinity levels (36 and 48 dS/m) up to 3.22 and 2.0 folds, respectively. Overall, S. aegyptiaca had a better potential of Na⁺ phytoremediation, and tolerated higher salinity compared to D. spicata. In contrast, the vigorous root and rhizome growth in D. spicata made it a proper solution for protecting the soils against further erosion under saline conditions.     

Modelling reveals endogenous osmotic adaptation of storage tissue water potential as an important driver determining different stem diameter variation patterns in the mangrove species Avicennia marina and Rhizophora stylosa

Background

Stem diameter variations are mainly determined by the radial water transport between xylem and storage tissues. This radial transport results from the water potential difference between these tissues, which is influenced by both hydraulic and carbon related processes. Measurements have shown that when subjected to the same environmental conditions, the co-occurring mangrove species Avicennia marina and Rhizophora stylosa unexpectedly show a totally different pattern in daily stem diameter variation.

Methods

Using in situ measurements of stem diameter variation, stem water potential and sap flow, a mechanistic flow and storage model based on the cohesion–tension theory was applied to assess the differences in osmotic storage water potential between Avicennia marina and Rhizophora stylosa.

Key results

Both species, subjected to the same environmental conditions, showed a resembling daily pattern in simulated osmotic storage water potential. However, the osmotic storage water potential of R. stylosa started to decrease slightly after that of A. marina in the morning and increased again slightly later in the evening. This small shift in osmotic storage water potential likely underlaid the marked differences in daily stem diameter variation pattern between the two species.

Conclusions

The results show that in addition to environmental dynamics, endogenous changes in the osmotic storage water potential must be taken into account in order to accurately predict stem diameter variations, and hence growth.     

Comparison of the effects of NaCl and KCl at the roots on seedling growth,cell death and the size,frequency and secretion rate of salt glands in leaves of <Emphasis Type="Italic">Limonium sinense</Emphasis>

Twenty days’ exposure to 50 or 100 mM NaCl in the rooting medium substantially increased fresh and dry weights of seedling shoots of the recretohalophyte Limonium sinense while 200 or 300 mM were increasingly inhibitory. KCl treatment was only slightly stimulating (50 mM) or strongly inhibitory (100–300 mM). Lesser effects on leaf area were also seen. Diameter of foliar salt glands was significantly larger than that of controls in 100 and 200 mM NaCl with the effect being reversed at higher concentrations. Gland enlargement was also observed in the presence of 100 mM KCl, while larger concentrations reduced gland size. Generally, gland diameter was larger in the presence of NaCl than in KCl. NaCl and KCl also increased gland number per leaf and secretion rate per gland. At 100 and 200 mM NaCl or KCl, Na⁺ secretion per leaf from NaCl-treated plants exceeded K⁺ secretion rate from KCl-treated plants while at 200 mM, Na⁺ secretion per gland was significantly higher for Na⁺ than for K⁺. Evidence of cell death in leaves of salt-treated plants using Evans blue staining indicates that release of cell contents through loss of membrane integrity contributed to the secretion values. We conclude that the greater tolerance of L. sinenseto to NaCl compared to KCl is linked to the more effective secretion of Na⁺ than of K⁺ and, in turn, to a greater stimulation of salt gland formation and activity and larger gland diameter.     

Ecophysiological and species‐specific responses to seasonal variations in halophytic species of the chenopodiaceae in a Mediterranean salt marsh

Plants adopt different strategies in response to seasonal variations in their natural biotype. In this study, the responses of seasonal changes on the organic osmolyte contents, ionic homoeostasis, osmotic potential and succulence of native chenopods in a salt marsh of Egypt were studied. Five halophytic species of the chenopodiaceae were collected during the rainy and dry season: Atriplex halimus L., Halocnemum strobilaceum (Pall.), Salicornia fruticosa (L.), Suaeda pruinosa (Lange) and Arthrocnemum macrostachyum (Moric). Results showed that plants employ an osmoconformer strategy via species specificity of response to seasonal variations. The total ions accumulation in all species was higher during the rainy season compared with the dry season. In particular, during rainy season, A. halimus resulted the higher accumulator species, followed by H. strobilaceum. Conversely, S. fruticosa, S. pruinosa and A. macrostachyum were defined as low‐accumulator species. In each studied species, the effective salinity, biological absorption coefficient as well as leaf succulence were higher during the rainy season. In conclusion, the present study have pointed out the importance that should be accorded with the development of research directed towards agronomic aspects for the uses of halophytic chenopodiaceae in salt marsh. Further, data obtained indicate that the ions accumulator species should be considered for use in phytoremediation of degraded saline lands.