Response of seagrass epiphyte loading to field manipulations of fertilization, gastropod grazing and leaf turnover rates

This study evaluates the bottom-up and top-down controls on epiphyte loads under low nutrient additions. Nutrients and gastropod grazers were manipulated in a field experiment conducted within a Thalassia testudinum meadow in Florida Bay, FL, USA. The effect of seagrass leaf turnover rate on epiphyte loading was also evaluated using novel seagrass short-shoot mimics that “grow,” allowing for the manipulation of leaf turnover rates. During the summer growing season and over the course of one seagrass leaf turnover period, low-level water column nutrient additions increased total epiphyte load, epiphyte chlorophyll a, and epiphyte autotrophic index. T. testudinum leaf nutrients (N and P) and leaf productivity also increased. Epiphyte loading and T. testudinum shoot biomass and productivity did not respond to a 60% mean increase in gastropod abundance. Manipulations of seagrass leaf turnover rates at minimum wintertime and maximum summertime rates resulted in a 20% difference in epiphyte loading. Despite elevated grazer abundances and increased leaf turnover rates, epiphyte loads increased with nutrient addition. These results emphasize the sensitivity of T. testudinum and associated epiphytes to low-level nutrient addition in a nutrient-limited environment such as Florida Bay.     

Biomass accumulation and shading effects of epiphytes on leaves of the seagrass, Heterozostera tasmanica, in Victoria,Australia

A method is described for estimating the rate of accumulation of epiphyte biomass on leaves of the seagrass, Heterozostera tasmanica (Martens ex Aschers.) den Hartog and for estimating the effect of epiphyte biomass on photosynthesis of the seagrass. Epiphyte biomass was determined by comparison of the weight per unit area of epiphyte-covered and epiphyte-free leaf blades. Epiphyte weight increased as age of the seagrass leaves increased. Linear regression on epiphyte biomass vs. leaf age estimated the rate of biomass accumulation. Rates varied from 5.7 to 104 μg epiphyte dry weight per cm² of leaf surface per day at three sites in Western Port and Port Phillip Bay, Victoria. Rates of accumulation of epiphyte biomass were generally higher during December through March (summer) than in May (autumn), August (winter) or October (Spring). Light attenuation by epiphytes increase linearly with biomass. The rate of biomass accumulation of epiphytes was compared with leaf growth rate, ambient photon flux density in H. tasmanica beds and the photosynthesis—photon flux density curve of H. tasmanica. This comparison demonstrated that epiphyte biomass can accumulate fast enough to shade H. tasmanica leaves and significantly reduce the time (to less than one half of the leaf life span) in which positive net photosynthesis of the leaf blade is possible.     

The short-term influence of herbivory near patch reefs varies between seagrass species

The coexistence of multiple species within a trophic level can be regulated by consumer preferences and nutrient supply, but the influence of these factors on the co-occurrence of seagrass species is not well understood. We examined the biomass and density responses of two seagrass species in the Florida Keys Reef Tract to grazing pressure near patch reefs, and evaluated how nutrient enrichment impacted herbivory dynamics. We transplanted Halodule wrightii (shoalgrass) sprigs into caged and uncaged plots in a Thalassia testudinum (turtlegrass) bed near a patch reef. Nutrients (N and P) were added to half of the experimental plots. We recorded changes in seagrass shoot density, and after three months, we measured above- and belowground biomass and tissue nutrient content of both species. Herbivory immediately and strongly impacted H. wrightii. Within six days of transplantation, herbivory reduced the density of uncaged H. wrightii by over 80%, resulting in a decrease in above- and belowground biomass of nearly an order of magnitude. T. testudinum shoot density and belowground biomass were not affected by herbivory, but aboveground biomass and leaf surface area were higher within cages, suggesting that although herbivory influenced both seagrass species, T. testudinum was more resistant to herbivory pressure than H. wrightii. Nutrient addition did not alter herbivory rates or the biomass of either species over the short-term duration of this study. In both species, nutrient addition had little effect on the tissue nutrient content of seagrass leaves, and N:P was near the 30:1 threshold that suggested a balance between N and P. The different impacts of grazing on these two seagrass species suggest that herbivory may be an important regulator of the distribution of multiple seagrass species near herbivore refuges like patch reefs in the Caribbean.     

A preliminary study of the seagrass Thalassodendron ciliatum (Forssk.) den hartog from Eastern Indonesia. Biological results of the snellius II expedition

Two dense meadows of the seagrass Thalassodendron ciliatum (Forssk.) den Hartog were sampled during the Indonesian—Dutch Snellius II expedition to Eastern Indonesia. Production data were obtained from one of these meadows. The production of leaf biomass was measured by the leaf marking technique of Zieman and by the plastochrone interval method. The two methods reached comparable results. The production of leaf tissue was 4.2 mg ADW shoot^?1 day^?1. The production of rhizome biomass was calculated in a similar way, based on the plastochrone interval of rhizome nodes. The production of the meadow, exclusive of the production of roots and fruits, amounted to 4.5 g ADW m^?2 day^?1. A significant correlation between the growth rates of rhizomes and leaves was observed. Biomass data from the second site are given.     

Fluorescence imaging application: effect of leaf age on seagrass photokinetics

We used the Imaging-PAM fluorometer to map spatial variability of photosynthesis in three seagrass species, Halophila ovalis, Zostera capricorni and Posidonia australis. Photosynthesis was described by relative photosynthetic rate (PS/50), effective quantum yield (Φ_PSII), non-photochemical quenching (NPQ and qN), electron transport rate (ETR) and leaf absorptivity. Photosynthetic patterns were linked to leaf age and light climate but patterns were not consistent across species. Longitudinal heterogeneity in photosynthesis was apparent along the leaves of all three species while lateral spatial heterogeneity was found only across Z. capricorni and H. ovalis leaves. Age of leaf tissue, determined by longitudinal location on the leaf, strongly influenced photosynthetic activity of Z. capricorni and P. australis. A comparison of H. ovalis leaves of differing maturity demonstrated the influence of leaf age on photosynthetic activity, yet a comparison of Z. capricorni leaves of differing maturity showed no leaf-age effects.Variations in stress-induced changes across a seagrass leaf can be used to identify areas or particular regions of the leaf, which are more susceptible to photodamage. Clear evidence of substantial within-leaf heterogeneity in photosynthetic activity (i.e., a two-fold variation in half saturation constant along a leaf of P. australis) has serious implications for use of small sections of leaf for photosynthetic incubations (such as O₂ or single-point chlorophyll a fluorescence measurements).     

<Emphasis Type="Italic">In vitro</Emphasis> collecting (IVC). I. The effect of collecting method and antimicrobial agents on contamination in temperate and tropical collections

Summary In vitro collecting is the process of initiating tissue cultures in the field. In order for in vitro collecting to be broadly available as a technique for collecting plant germplasm, the levels of contamination in such cultures must be controlled. Two techniques for in vitro collecting were compared: leaf punch and needle collecting. The effectiveness of these methods for collecting leaf and stem tissues from plants at tropical and temperature sites was compared. Stem tissue collected by the needle collecting method gave cultures with an average contamination percentage of 31% and 16%, from the tropical and temperate sites, respectively, while with the leaf punch method, average contamination percentages were 90% and 69%. The effectiveness of antimicrobial agents in reducing contamination in leaf punch cultures was evaluated. Addition of the fungicide benlate and the antibiotics cefotaxime and vancomycin, to the leaf punch collections reduced contamination to an average of 30% in the tropical collections and 35% in the temperate collections. Over 90% of both tropical and temperature species collected in multiple samples of 10 or more had at least one clean sample using this medium. The use of either the leaf punch method in combination with a fungicide and antibiotics or the needle collecting technique yielded a high percentage of clean tissues for study and growth.     

The photosynthetic performance of the tropical seagrass Halophila ovalis in the upper intertidal

Some ecophysiological adaptation strategies of the tropical seagrass Halophila ovalis were investigated with respect to this plant's ability to grow in the upper intertidal in either monospecific pools (but not together with other intertidal species) or emergent and exposed to high temperatures and irradiances during several hours every day. It was found that Halophila ovalis could raise the pH in simulated pools to 8.6, while the two other major (biomass wise) intertidal seagrasses raised the pH to 8.8 (Cymodocea rotundata) and 9.2 (Thalassia hemprichii). In situ, midday pH values of 8.5, 8.7 and 9.0 were recorded in pools inhabiting the three species, respectively. It was further found that photosynthetic electron transport rates (ETR) measured continuously in situ for one leaf during a diurnal cycle followed the irradiance up to a daily maximal value of 2200 μmol photons m⁻² s⁻¹, but dropped by ca. 50% when the leaf became air exposed just before noon. This drop in ETR upon emergence was verified by “point measurements”, while ETRs of leaves that grew in small pools formed as the tide receded followed the irradiance more closely. Similarly, F_v/F_m measured after 15 min of dark adaptation decreased by ca. 50% in emergent leaves during midday, but maintained higher daily values in the submerged leaves.     

The role of leaf nitrogen content in determining turtlegrass (Thalassia testudinum) grazing by a generalized herbivore in the northeastern Gulf of Mexico

In shallow marine environments the variability in grazing on seagrasses has been hypothesized to be controlled, in part, by the nutritive quality (i.e., nitrogen content) of their leaves. The few existing studies of the relationship between leaf nitrogen content and seagrass grazing have all found a positive relationship between leaf nitrogen content and preference by selective vertebrate grazers (i.e., the bucktooth parrotfish, green sea turtles, and dugongs). However, most marine herbivores (both vertebrate and invertebrate) are thought to be extreme generalists with broad diets of variable nutritive quality (e.g., detritus, living plants, and animals), suggesting the currently held view on the role leaf nutrient content in explaining the variability of seagrass grazing is an oversimplification.In this study, we evaluated how leaf nitrogen content influenced grazing on turtlegrass by a generalist invertebrate herbivore (the pink sea urchin Lytechinus variegatus) in the northeastern Gulf of Mexico. Using a short-term laboratory test and a longer-term field experiment, we tested the hypothesis that leaf nitrogen content controls sea-urchin grazing on seagrass leaves. We hypothesized that if poor nutritive value of seagrasses is responsible for reduced rates of feeding, then increasing leaf nitrogen concentrations should lead to increased rates of seagrass consumption by sea urchins.In the field experiment, we significantly enriched seagrass leaf nitrogen concentrations (some 10-20% depending on month) in experimental plots with a commercial fertilizer and we manipulated grazing intensity by enclosing adult sea urchins at densities that bracketed the range of average densities observed in the region (i.e., 0, 10 and 20 individuals/m(2)). Comparisons of changes in aboveground seagrass production and biomass showed no evidence that sea urchins grazed significantly more in treatments where leaf nitrogen was enriched. Because the statistical power of our test to detect such differences was low and aboveground seagrass production varied significantly among treatments, we also used a mass balance equation to estimate sea urchin consumption of nitrogen-enriched and unenriched leaves. This showed that sea urchins compensated for low nitrogen levels in our unenriched treatments by eating more leaves than in treatments where leaf nitrogen was elevated. Using a laboratory test, we also found that sea urchins ate less nitrogen-enriched seagrass than unenriched seagrass. In combination, these results show that, in contrast to findings reported for vertebrate herbivores, sea urchins feed at higher rates when offered seagrass leaves of lower leaf nitrogen content, and that low levels of leaf nitrogen are not always an effective defense against herbivores.     

Distribution and leaf growth of Thalassodendron pachyrhizum den Hartog in Southern Western Australia

The seagrass Thalassodendron pachyrhizum den Hartog grows on limestone reef platforms. Monthly leaf biomass was measured over 2 years and showed a strong seasonal variation with maximum biomass of 500 g m⁻². This seagrass loses all its leaves except for a bud and this characteristic was used to obtain a conservative estimate of productivity by change in standing stock. Leaf growth during the growing season was 6.6 mg Cg⁻¹ day⁻¹. Leaf length frequencies showed that new leaves formed during autumn (March–April). They grew from autumn until spring (November) and began to senesce in summer, followed by leaf fall in late summer (February–March).The growth of rhizome shoots “invading” free substratum space and the growth of new stems was measured for a 300-day period; about 9 leaves were produced in this period.     

Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

Correlations between salinity and growth of the seagrass Amphibolis antarctica (labill.) Sonder & Aschers., In Shark Bay,Western Australia,using a new method for measuring production rate

A method of estimating above-ground productivity in situ of the seagrass Amphibolis antarctica (Labill.) Sonder & Aschers. has been devised, using tags to determine rates of leaf turnover. This has proved an effective tool in establishing the behavior of the species in relation to the gradient of increasing salinity which is present in Shark Bay. No seagrass was found beyond 64%_o, but measurement of production and biomass within dense patches of seagrass at different salinities revealed that these were at a maximum at a salinity of 42%_o, decreasing as the salinity increased and also at lower oceanic concentrations. Production rates ranged from 2 to 17 g dry weight m^?2 day^?1 with biomass from 600 to 2000 g m^?2, thus Amphibolis antarctica is one of the more productive Australian seagrass species, even in the hypersaline conditions of the Bay. Despite the obvious correlation between above-ground production and salinity, it is pointed out that the results are not taken to imply causality.     

Do desiccation tolerances control the vertical distribution of intertidal seagrasses?

Photosynthetic processes in Zostera japonica, an upper intertidal species, were found to be more severely affected by desiccation than Z. marina, a lower intertidal and subtidal species, at comparable levels of tissue water content. The data indicate that photosynthetic responses to desiccation at the level of the individual leaf are insufficient to explain observed patterns of intertidal seagrass zonation. Desiccation tolerance in seagrasses is more likely to involve a complex interaction of morphological traits and growth strategies at the level of the whole plant, such as downsizing (e.g. smaller, narrower leaves), reduced structural rigidity and increased rates of leaf abscission and leaf turnover.     

Variations in sensitivity to copper and zinc among three isolated populations of the seagrass, Zostera capricorni

Metal accumulation in seagrass is well documented, but toxic impacts and mechanisms of tolerance in seagrass are not well understood. We looked at the impacts of 10 h exposure to copper and zinc for three isolated populations of Zostera capricorni in the Sydney (Australia) region. Photosynthetic efficiency (measured as the effective quantum yield, ΔF/Fm′) and chlorophyll pigment concentrations showed different sensitivities to metal impacts at the three geographically isolated sites. Seagrasses from the least developed estuary were the most sensitive to metals and the two more developed estuaries had more tolerant populations. Determination of metal concentrations in the leaves showed that there was no difference in metal exclusion as the sensitive seagrass accumulated no more metal than the tolerant seagrass. Equally, background levels of copper and zinc in the sediments and seagrass tissue could not explain the differences in tolerance. We discuss some other possible mechanisms of tolerance. The outcomes suggest that assessing metal content in seagrass tissue may not demonstrate degree of photosynthetic impact.     

Grazing effects of a gammaridean Amphipoda,Ampithoe sp., on the seagrass,Syringodium isoetifolium,and epiphytes in a tropical seagrass bed of Fiji

In the pure stand of tropical seagrass,Syringodium isoetifolium, in a small oceanic island, Fiji, grazing effects of the seagrass-associated gammarid,Ampithoe sp., on seagrass and epiphytes were assessed in October 1989, November 1991, November 1992. Density of the gammarid was estimated with two methods, mesh bag method and tuft method. During the three years surveyed the density of the gammarid increased remarkably from 1989 to 1991, with heavy epiphytism. Gut contents of the gammarid were examined. Grazing rates on seagrass leaf with and without epiphytic blue-green algae were measured in a bottle experiment. Litter bag experiments were conducted using different mesh sizes each containing seagrass only and seagrass and gammarids. The seagrass leaf biomass in the litter bag reduced abruptly in both bags. After one week, 78–86% of seagrass biomass disappeared from the bags. Enhancement of decomposition of seagrass leaf by the gammarid grazing was observed. Oxygen consumption and ammonium excretion rates were measured simultaneously in bottle experiments. Carbon budget in the seagrass bed was estimated as follows: 0.9 gC m⁻² day⁻¹ in seagrass growth, gammarid grazing was about a half of it and further assimilated a half of it, about 0.1 gC m⁻² day⁻¹, and more than half of it become CO₂ by respiration. Grazing effects on epiphyte and seagrass growth and production were discussed through the carbon budget and indirect interactions between seagrass, epiphytes and associated gammarids to explain the temporal change of seagrass and epiphyte dynamics.     

Heterophylly results in a variety of “spectral signatures” in aquatic plant species

The leaf reflectance spectra (280–887 nm) of two heterophyllous aquatic plant species Polygonum amphibium (L.) and Nuphar luteum (L.) were compared and their relation to physical properties of the leaves examined. In P. amphibium contrasting environmental conditions along water–land gradient affected the majority of anatomical and morphological properties of leaves, but less differences were observed in photosynthetic pigment and total flavonoid contents. Leaf mass per area (LMA), palisade mesophyll, leaf thickness, trichome length and anthocyanin content per dry mass were correlated to the different parts of spectra. In N. luteum natant and submerged leaves differed significantly in all measured parameters. Chlorophyll a, anthocyanin and carotenoid contents per dry mass were related to reflectance in the red region, while leaf thickness, anthocyanin and total flavonoid contents per leaf area were related to reflectance in the near infrared region. Redundancy Analysis (RDA) indicated that in P. amphibium the average length of trichomes and LMA explained 72% and 6% variability of the spectra, whereas in N. luteum anthocyanin content per dry mass, explained 57% variability of the spectra. The comparison of natant leaves of both species showed that they were more similar than different leaf types within the single species.     

A comparative analysis of leaf shape of wheat, barley and maize using an empirical shape model

Background and Aims

The phenotypes of grasses show differences depending on growth conditions and ontogenetic stage. Understanding these responses and finding suitable mathematical formalizations are an essential part of the development of plant and crop models. Usually, a marked change in architecture between juvenile and adult plants is observed, where dimension and shape of leaves are likely to change. In this paper, the plasticity of leaf shape is analysed according to growth conditions and ontogeny.

Methods

Leaf shape of Triticum aestivum, Hordeum vulgare and Zea mays cultivars grown under varying conditions was measured using digital image processing. An empirical leaf shape model was fitted to measured shape data of single leaves. Obtained values of model parameters were used to analyse the patterns in leaf shape.

Key Results

The model was able to delineate leaf shape of all studied species. The model error was small. Differences in leaf shape between juvenile and adult leaves in T. aestivum and H. vulgare were observed. Varying growth conditions impacted leaf dimensions but did not impact leaf shape of the respective species.

Conclusions

Leaf shape of the studied T. aestivum and H. vulgare cultivars was remarkably stable for a comparable ontogenetic stage (leaf rank), but differed between stages. Along with other aspects of grass architecture, leaf shape changed during the transition from juvenile to adult growth phase. Model-based analysis of leaf shape is a method to investigate these differences. Presented results can be integrated into architectural models of plant development to delineate leaf shape for different species, cultivars and environmental conditions.     

Leaf dimorphism in Thuja plicata and Platycladus orientalis (thujoid Cupressaceae s. str., Coniferales): the changes in morphology and anatomy from juvenile needle leaves to mature scale leaves

Thuja plicata and Platycladus orientalis initially produce only bifacial needle leaves. When the first lateral shoots develop, the leaf morphology and anatomy changes dramatically. Subsequently, only greatly reduced, bifacial scale leaves are developed. A new kind of “superimposed bifaciality” occurs with the change from juvenile needle leaves to mature scale leaves. Anatomical dorsiventrality affects not only the individual leaf, but also the complete plagiotropic lateral shoots of Thuja, which have a sun- and shade-exposed side. The upper light-exposed median leaves show adaxial leaf anatomy, contrary to the lower shaded median leaves showing abaxial leaf anatomy. Due to their mixed exposure, the lateral leaves show a lateral differentiation. At vertical lateral shoots of Platycladus, a predominant light-exposed side is absent. Thus, the anatomical dorsiventrality does not affect the complete shoot. Here the morphological abaxial side of a scale leaf becomes functionally and physiologically adaxial by reorientation of the palisade parenchyma and stomata. In juvenile needle leaves, the palisade parenchyma is located adaxial, with the majority of stomata being located abaxial. Conversely, in mature scale leaves, the palisade parenchyma is abaxial and the majority of stomata are adaxial.     

The seasonal distribution and community structure of the epiphytic algae on Thalassia hemprichii (Ehrenb.) Aschers. from Papua New Guinea

Algae growing as epiphytes on leaves of Thalassia hemprichii (Ehrenb.) Aschers. have been studied from November 1980 to December 1981, in the Port Moresby area, Papua New Guinea. The epiphytic communities of 3 different monospecific seagrass meadows are compared for species richness, abundance and temporal pattern. Seagrass shoots were studied separately, using the method of Braun—Blanquet, as adapted by Boudouresque. By differentiating between the leaves of one single shoot, the inner- and outer-face of each leaf and the upper- and lower-part of each leaf, the epiphytic community was studied from its initial colonization (Leaf 1) to the final “climax” situation (Leaf 4). The diversity and abundance were strongly related to the age of the seagrass leaves. The Rhodophyta were best represented, with the Cryptonemiales dominating the community quantitatively; the Ceramiales predominated qualitatively. The Phaeophyta were negligible in terms of abundance and diversity. Differences between the 3 study sites are presented.     

The ethnobotany and pharmacognosy of Olea europaea subsp. africana (Oleaceae)

The ethnobotanical uses of wild olive, O. europaea subsp. africana (sometimes referred to as subsp. cuspidata) in southern Africa and in other parts of Africa are reviewed. Chromatographic analyses of secoiridoids (oleuropein and other oleuropeosides) in 25 wild olive leaf samples from 10 localities in South Africa showed substantial amounts of oleuropein (up to 110 mg/g dry weight) and not trace amounts as reported in the literature. Oleuropein is the main active compound in olive leaf, with demonstrated anti-oxidant, anti-microbial, hypolipidemic and hypotensive activities. A comparison with nine cultivated olive leaf samples (subsp. europaea) from six cultivars and two localities showed that commercial olive leaf can be distinguished by the presence of verbascoside, which is absent in wild olive. Extraction methods and solvent systems (TLC and HPLC) were compared, using pure oleuropein (isolated from wild olive leaf and identified by NMR) as an authentic reference sample. The unique peltate scales on the leaves are useful to identify olive leaf raw material (but are the same in both subspecies). The main conclusion is that wild olive leaf is chemically closely similar to cultivated olive leaf and therefore suitable as an alternative source of raw material for olive leaf extract.