Production dynamics of the eelgrass, <Emphasis Type="Italic">Zostera marina</Emphasis> in two bay systems on the south coast of the Korean peninsula

Production dynamics of eelgrass, Zostera marina was examined in two bay systems (Koje Bay and Kosung Bay) on the south coast of the Korean peninsula, where few seagrass studies have been conducted. Dramatically reduced eelgrass biomass and growth have been observed during summer period on the coast of Korea, and we hypothesized that the summer growth reduction is due to increased water temperature and/or reduced light and nutrient availabilities. Shoot density, biomass, morphological characteristics, leaf productivities, and tissue nutrient content of eelgrass were measured monthly from June 2001 to April 2003. Water column and sediment nutrient concentrations were also measured monthly, and water temperature and underwater irradiance were monitored continuously at seagrass canopy level. Eelgrass shoot density, biomass, and leaf productivities exhibited clear seasonal variations, which were strongly correlated with water temperature. Optimal water temperature for eelgrass growth in the present study sites was about 15–20°C during spring period, and eelgrass growths were inhibited at the water temperature above 20°C during summer. Daily maximum underwater photon flux density in the study sites was usually much higher than the light saturation point of Z. marina previously reported. Densities of each terminal, lateral, and reproductive shoot showed their unique seasonal peak. Seasonal trends of shoot densities suggest that new eelgrass shoots were created through formation of lateral shoots during spring and a part of the vegetative shoots was transformed into flowering shoots from March. Senescent reproductive shoots were detached around June, and contributed to reductions of shoot density and biomass during summer period. Ambient nutrient level appeared to provide an adequate reserve of nutrient for eelgrass growth throughout the experimental period. The relationships between eelgrass growth and water temperature suggested that rapid reductions of eelgrass biomass and growth during summer period on the south coast of the Korean peninsula were caused by high temperature inhibition effects on eelgrass growth during this season.     

Leaf reddening in the seagrass Thalassia testudinum in relation to anthocyanins, seagrass physiology and morphology, and plant protection

Numerous seagrass species growing in high-light environments develop red coloration in otherwise green leaves, yet the ecophysiology of leaf reddening in seagrasses is poorly understood. To increase our understanding of the process of leaf reddening in Thalassia testudinum found in the lower Florida Keys (USA), we identified the molecules responsible for red coloration in leaves and compared physiological, morphological, and growth attributes of entirely red-leafed shoots to entirely green-leafed shoots. We determined that four anthocyanin molecules are responsible for red coloration in leaves. In addition, we found that red leaves had higher concentrations of photoprotective pigments (anthocyanins and UV-absorbing compounds), higher effective quantum yields (ΔF/F _m′) at midday, and were shorter, narrower, and weighed less than green leaves. No significant difference in growth rates was observed between red- and green-leafed shoots, but patches of red-leafed shoots had shorter canopy heights and smaller LAI compared to patches of green-leafed shoots. Our results demonstrate that leaf reddening in T. testudinum is caused by high concentrations of anthocyanins, is associated with physiological and morphological attributes, and acts as a sunscreen since red leaves were able to maintain high effective quantum yields at high light intensities.     

Eelgrass fairy rings: sulfide as inhibiting agent

Distinct ‘fairy rings’ consisting of narrow fringes of eelgrass (Zostera marina L.) expand radially over a bottom of chalk plates outside the calcium carbonate cliffs of the island of Møn, Denmark. We conducted a survey to evaluate possible explanations for the formation of the rings and, more specifically, for the apparent die-off of eelgrass shoots on the inner side of the rings. The fairy rings were up to 15 m in diameter consisting of 0.3- to 1-m-wide zones of sea grass shoots at densities of up to 1,200 shoots m^?2 and rooted in an up to 10-cm-thick sediment layer. On the outer side, shoots expanded over the bare chalk plates. On the inner side, shoots were smaller, had lower absolute and specific leaf growth, shoot density was lower and the sediment eroded leaving the bare chalk with scattered boulders behind. Sediment organic matter and nutrients and tissue nutrient contents were not different among positions. Sediment pools of acid volatile sulfides and chromium-reducible sulfur increased from outer to the middle positions of the rings, and so did total sulfur content of eelgrass tissues, while tissue δ³⁴S isotope ratios, regardless of position in the fringes, were low reflecting substantial invasion of sulfide from the sediment. Neither the clonal growth pattern of eelgrass, sediment burial of shoots, hydrodynamic forcing nor nutrient limitation could explain the ring-shaped pattern. We conclude that the most likely explanation must be found in invasion of eelgrass shoots by toxic sulfide accumulating in the sediment due to low iron availability in the carbonate-rich environment.     

Influence of sediment nitrogen-availability on carbon and nitrogen dynamics in the seagrass Thalassia testudinum

Variations in tissue carbon (C), nitrogen (N) content, and non-structural carbohydrate (NSC) reserves in the turtle grass Thalassia testudinum Banks ex König were examined in relation to changes in sediment-N availability in Corpus Christi Bay (CCB) and lower Laguna Madre (LLM), Texas, USA, from May to October 1997. Under natural conditions, sediment pore-water NH⁺ ₄-concentrations were higher at CCB (100?μM) than at LLM (30?μM); this difference was reflected in a significantly higher leaf and rhizome N-content at CCB than at LLM. However, sediment NH⁺ ₄-enrichment using a commercial fertilizer resulted in significantly higher tissue N-content relative to controls at both sites. N enrichment also influenced plant carbon metabolism, as reflected by distinct increases in leaf C-content at both sites. Significant decreases in rhizome NSC-content was recorded during the first two months of the experiment, suggesting that C was reallocated from rhizomes to leaves to support stimulated leaf growth at both sites. At LLM, leaf growth-rates increased and leaf turnover-time decreased as a result of sediment NH⁺ ₄-enrichment. With respect to chlorophyll, concentrations did not change significantly at CCB, but increased steadily at LLM after the first month following fertilization. In general however, chlorophyll concentrations in control plots were significantly higher at CCB than that at LLM. These observations suggest that leaf function related to C-fixation is enhanced under higher sediment N conditions, as reflected in higher leaf growth-rates and increased blade chlorophyll-content. In contrast, under low-N conditions, below-ground tissue production is enhanced at the expense of the above-ground shoots and leaves, resulting in the high below: above-ground biomass ratios often observed in seagrass beds of oligotrophic environments.     

Theoretical analysis of change in forest carbon use efficiency with stand development: A case study on Hinoki Cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) plantation from the seedling stage

Changes in carbon use efficiency (CUE), which is defined as the ratio of net primary production (NPP) to gross primary production (GPP), were estimated for the aerial parts of the Hinoki Cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) with respect to stand development. The analysis incorporated previously published data from the early stages of stand development, namely the seedling stages of the cypress. For this analysis, a simple mathematical model to assess the changes in CUE was developed by incorporating data on physiological variables and mass of woody species. The CUE tended to increase with increases in the aboveground biomass of the stand, and then decreased gradually despite increases in the aboveground biomass. The CUE-value (0.28, 0.39) of the seedling stage was lower than that (0.33-0.58) of the young or mature trees. To examine the effect of physiological variables and mass on CUE, the ratios of the specific respiration rate to the specific photosynthetic rate (r/a) and the leaf biomass to the aboveground biomass or leaf mass ratio (y_L/y_T) were calculated. The low value of CUE at the seedling stage was due to the high ratio of specific respiration rate to specific photosynthetic rate r/a, but was not due to the high value of the leaf mass ratio y_L/y_T. In addition, the decline in CUE associated with older stages of stand development was due to the decreasing changes in y_L/y_T, and the r/a ratio did not influence the change in CUE.     

Mechanisms of moisture stress in a mid-latitude temperate forest: Implications for feedforward and feedback controls from an irrigation experiment

While it is well established that stomata close during moisture stress, strong correlations among environmental (e.g., vapor pressure deficit, soil moisture, air temperature, radiation) and internal (e.g., leaf water potential, sap flow, root-shoot signaling) variables obscure the identification of causal mechanisms from field experiments. Models of stomatal control fitted to field data therefore suffer from ambiguous parameter identification, with multiple acceptable (i.e., nearly optimal) model structures emphasizing different moisture status indicators and different processes. In an effort to minimize these correlations and improve parameter and process identification, we conducted an irrigation experiment on red maples (Acer rubrum L.) at Harvard Forest (summers of 2005 and 2006). Control and irrigated trees experienced similar radiative and boundary layer forcings, but different soil moisture status, and thus presumably different diurnal cycles of internal leaf water potential. Measured soil moisture and atmospheric forcing were used to drive a transient tree hydraulic model that incorporated a Jarvis-type leaf conductance in a Penman–Monteith framework with a Cowan-type (resistance and capacitance) tree hydraulic representation. The leaf conductance model included dependence on both leaf matric potential, Ψ_L (so-called feedback control) and on vapor pressure deficit, D (so-called feedforward control). Model parameters were estimated by minimizing the error between predicted and measured sap flow. The whole-tree irrigation treatment had the effect of elevating measured transpiration during summer dry-downs, demonstrating the limiting effect that subsurface resistance may have on transpiration during these times of moisture stress. From the best fitted model, we infer that during dry downs, moisture stress manifests itself in an increase of soil resistance with a resulting decrease in Ψ_L, leading to both feedforward and feedback controls in the control trees, but only feedforward control for the irrigated set. Increases in the sum-of-squares error when individual model components were disabled allow us to reject the following three null hypotheses: (1) the f(D) stress is statistically insignificant (p = 0.01); (2) the f(Ψ_L) stress is statistically insignificant (p = 0.07); and (3) plant storage capacitance is independent of moisture status (p = 0.07).     

Inhibition of the growth of micro-algae and bacteria by extracts of eelgrass (Zostera marina) leaves

The effects of the water-soluble fraction of dead leaves of the eelgrass Zostera marina L. on the growth of 8 species of micro-algae (pennate and centric diatoms, dinoflagellates, and a green flagellate) and a bacterium were studied on agar plates and in liquid culture. The extracts of leaves which had been dead from a few days to 2 wk inhibited growth and often killed cells in all test organisms. Extracts were lethal even at concentrations equivalent to as little as 0.25 mg dry leaf ml^-1, but inhibition decreased when extracts were prepared from leaves aged in the laboratory for 35 d (loss of anti-bacterial activity) or 90 d (loss of anti-algal activity). Extracts of leaves which had aged and dried several months in the field had no effect, except at very high concentrations (13 mg dry leaf ml^-1) when the lag phase of growth was prolonged several days in a culture of the chlorophyte Platymonas sp. The active fraction in eelgrass leaves may be important in controlling initial growth of micro-organisms on eelgrass detritus, and it could determine the composition and activity of the epiphytic community on living leaves.     

Significance of salinity and silicon levels for growth of a formerly estuarine eelgrass (Zostera marina) population (Lake Grevelingen, The Netherlands)

Since the early 1980s, the eelgrass, Zostera marina L., population in the saline Lake Gevelingen, The Netherlands, is rapidly declining. An earlier study, in which long-term data on eelgrass coverage in this former estuary were correlated with several environmental variables, showed only one significant correlation: coverage was positively related to water column silicon levels. In addition, a negative correlation with salinity was observed, but this was not significant. In the present study, the effect of silicon and the effect of salinity on the development of Z. marina were investigated experimentally. Enhancement of dissolved silicon concentrations in the water did not stimulate Z. marina above-ground production or an increase in final above- and below-ground biomass. The highly significant correlation between eelgrass coverage and water column silicon levels, thus, remains to be explained. The results of the growth experiments did, however, demonstrate a clear effect of salinity on Z. marina growth. Plants cultured at 22 psu showed a higher production of shoots and leaves, resulting in more above-ground biomass, than plants grown at 32 psu. In addition, below-ground biomass was also higher at 22 psu. Measurements of chlorophyll a fluorescence, performed with a PAM-fluorometer, indicated a reduction of photosynthesis in the high-salinity treatments. Thus, low salinity stimulates development of Z. marina from Lake Grevelingen. Eelgrass from such a historically estuarine area may be more sensitive to high salinities than other, more marine populations. Recovery of the autochthonous eelgrass population is expected to be favoured when the estuarine conditions of the seagrass area are re-established, or when restoration programmes are carried out with allochthonous ecotypes that are less sensitive to high salinities. Received: 23 June 1998 / Accepted: 19 November 1998     

Comparisons in demographic rates of bay scallops in eelgrass and the introduced alga, Codium fragile, in New York

Bay scallops, Argopecten irradians, supported vibrant fisheries which subsequently collapsed, as such, they are a focus species for many restoration efforts along the Atlantic and Gulf coasts of the United States. The scallops’ preferred habitat, seagrass, has also dramatically declined, and some scallop populations have increased post-restoration despite reduced seagrass cover. This has led to the hypothesis that macroalgae may serve as suitable alternative habitats for bay scallops. This study is the first to compare demographic rates, such as long-term survival, growth, condition and reproductive potential of scallops between the native eelgrass, Zostera marina, and the introduced alga, Codium fragile. Although long-term survival was not different between habitats, results suggest site-specific and inter-annual variation in the impacts of Codium on scallop growth. While demographic rates did not differ in Shinnecock Bay, in Sag Harbor, growth and/or condition were significantly different between both vegetated habitats depending on the year. However, recruit density, size and condition did not vary significantly, adding to the complexity of this relationship. Despite potential site-specific and inter-annual differences, this study supports the hypothesis that habitats other than eelgrass can benefit bay scallops.     

Leaf-root interaction in the uptake of ammonia by Zostera marina

The effect of ammonia uptake by one organ on the uptake of that nutrient by another organ was determined for the seagrass Zostera marina L. under laboratory conditions. Leafy shoots with roots attached were incubated in two-compartment chambers, with the roots in one compartment and the leaves in the other. Rates of ammonia uptake were compared when roots and leaves were supplied with ammonia separately and simultaneously. Root uptake of ammonia had no influence on the rate of ammonia uptake by leaves. However, leaf uptake of ammonia caused a 77% decrease in the maximum rate of ammonia uptake by roots. The K_s values for ammonia uptake by leaves and roots were 9.2 and 104 M respectively, showing that leaves had a greater affinity for ammonia than roots.     

Drastic decline of an extensive eelgrass bed in Nova Scotia due to the activity of the invasive green crab (Carcinus maenas)

In 2001–2002, there were severe declines of eelgrass (Zostera marina) in estuaries along the Nova Scotia coast of the southern Gulf of St. Lawrence. We examined the relationship between the recent invasion of European green crabs (Carcinus maenas) and the eelgrass decline in Benoit Cove in Tracadie Harbour. The abnormally abundant eelgrass wrack consisted mainly of entire shoots, not the usual blades. Three separate methods yielded similar estimates of the rate of shoot removal from the eelgrass beds (direct quadrat counts, “mark–recapture” of tagged shoots in the eelgrass bed, and rate of shoot appearance in the shore wrack). From 14 July to 8 September 2002, the shoot density in the bed fell by about 75 %. Similar rates of decline occurred in crab enclosures (4.4 crabs m^?2) placed in the eelgrass bed. Green crab foraging, involving the tearing of shoots and the digging of large pits, was the reason for the drastic decline of the eelgrass bed in Benoit Cove. It is clear that an invasion of green crabs to a region where they had not previously existed can both destroy eelgrass beds and restrict their recovery.     

Mathematical analysis of change in forest carbon use efficiency with stand development: A case study on Abies veitchii Lindl.

Changes in carbon use efficiency (CUE), which is defined as the ratio of net primary production (NPP) to gross primary production (GPP), were analyzed for Abies veitchii Lindl. forests with respect to stand development by developing a simple mathematical model incorporating data on physiological variables and leaf mass ratio. A decrease in CUE with stand development was successfully expressed as a function of stand biomass (y) based on the following three assumptions: (1) a power-law relationship between mean respiration and mean individual tree mass, (2) a power-functional relationship between mean gross primary production and mean individual tree mass, and (3) self-thinning relationship between stand biomass and density. Based on this model, a parameter of CUE–y relationship was defined, and it was clarified that CUE decrease with stand development is caused not by the ratio of specific respiration rate to specific gross photosynthetic rate, but by leaf mass ratio. Since CUE is high in young forests, helpful information on selecting woody species when planting seedlings was provided from the viewpoints of reducing CO₂ in the atmosphere and global warming.     

Application of a temperature-dependent von Bertalanffy growth model to bullhead (Cottus gobio)

The most studied and commonly applied model of fish growth is the von Bertalanffy model. However, this model does not take water temperature into account, which is one of the most important environmental factors affecting the life cycle of fish, as many physiological processes that determine growth, e.g. metabolic rate and oxygen supply, are directly influenced by temperature. In the present study we propose a version of the von Bertalanffy growth model that includes mean annual water temperatures by correlating the growth coefficient, k, explicitly and the asymptotic length, L_∞, implicitly to water temperature. All relationships include parameters with an obvious biological relevance that makes them easier to identify. The model is used to fit growth data of bullhead (Cottus gobio) at different locations in the Bez River network (Drme, France). We show that temperature explains much of the growth variability at the different sampling sites of the network.     

Effects of in situ CO2 enrichment on the structural and chemical characteristics of the seagrass Thalassia testudinum

Seagrasses commonly display carbon-limited photosynthetic rates. Thus, increases in atmospheric pCO₂, and consequentially oceanic CO_2(aq) concentrations, may prove beneficial. While addressed in mesocosms, these hypotheses have not been tested in the field with manipulative experimentation. This study examines the effects of in situ CO_2(aq) enrichment on the structural and chemical characteristics of the tropical seagrass, Thalassia testudinum. CO_2(aq) availability was manipulated for 6 months in clear, open-top chambers within a shallow seagrass meadow in the Florida Keys (USA), reproducing forecasts for the year 2100. Structural characteristics (leaf area, leaf growth, shoot mass, and shoot density) were unresponsive to CO_2(aq) enrichment. However, leaf nitrogen and phosphorus content declined on average by 11 and 21 %, respectively. Belowground, non-structural carbohydrates increased by 29 %. These results indicate that increased CO_2(aq) availability may primarily alter the chemical composition of seagrasses, influencing both the nutrient status and resilience of these systems.     

Analysis of annual fluctuations of C. nodosa in the Venice lagoon: Modeling approach

A simple simulation model was developed to describe the growth trends of Cymodocea nodosa (Ucria) Ascherson based on data sets from the Venice lagoon. The model reproduces the seasonal fluctuations in the above and belowground biomass and in shoot density. The modeling results are in good agreement with data on net production, growth rates and chemical–physical parameters of water. It was assumed that light and temperature are the most important factors controlling C. nodosa development, and that the growth was not limited by nutrient availability. The aim was to simulate biomass production as a function of external forcing variables (light, water temperature) and internal control (plant density). A series of simulation experiments were performed with the basic model showing that among the most important phenomena affecting C. nodosa growth are: (1) inhibition of production and recruitment of new shoots by high temperature and (2) light attenuation due to seasonal fluctuation.     

The role of isopods and amphipods in the initial fragmentation of eelgrass detritus in Nova Scotia,Canada

Daytime observations on the isopods Idotea phosphorea and I. baltica and the amphipod Gammarus oceanicus held in laboratory microcosms showed that I. phosphorea and G. oceanicus spent 45% and 30% respectively, of their active time feeding on dead, intact eelgrass leaves which had been recently released from plants. I. baltica spent 41% of its active time consuming intact green leaves. The shredding of intact dead leaves by I. phosphorea and G. oceanicus resulted in production of small detrital particles which were liberated from the faeces of the invertebrates and this type of feeding led to the breakdown of whole leaves. Field experiments which separated the effects of shredding by invertebrates and grinding by waves and ice on the loss of weight from leaf packs showed that relative to controls isopods significantly increased weight loss from dead leaves. Loss of weight from leaf packs exposed to both biotic and physical shredding forces was not significantly different from that found on those exposed only to shredding by isopods. However, trends in the data indicated that fragmentation of whole, dead leaves in the field probably is a result of the synergistic effects of shredding by invertebrates and physical factors, particularly ice grinding. The role played by invertebrates in fragmenting intact, dead leaves is discussed in the light of energy flow and nutrient cycling within seagrass systems.     

Integrating effects of leaf nitrogen,age, rank,and growth temperature into the photosynthesis-stomatal conductance model LEAFC3-N parameterised for barley (Hordeum vulgare L.)

A crucial challenge for including biophysical photosynthesis–transpiration models into complex crop growth models is to integrate the plasticity of photosynthetic processes that is related to factors like nitrogen (N) content, age, and rank of leaves, or to the adaptation of plants to growth temperature (T_g). Here we present a new version of the combined photosynthesis-stomatal conductance model LEAFC3-N [Müller, J., Wernecke, P., Diepenbrock, W., 2005. LEAFC3-N: a nitrogen sensitive extension of the CO₂ and H₂O gas exchange model LEAFC3 parameterised and tested for winter wheat (Triticum aestivum L.). Ecological Modelling 183, 183–210.] that was revised, extended and completely re-parameterised for barley (Hordeum vulgare L.) with special regard for these factors to facilitate the use of the model in ecophysiological studies and in crop modelling. The analysis is based on novel comprehensive data on photosynthetic CO₂ and light response curves measured at two oxygen concentrations and different temperatures on leaves of barley (H. vulgare L.) differing in leaf N and chlorophyll content. Plants were grown in climatic chambers or in the field at different N and T_g.We thoroughly revised the existing and introduced new nitrogen relations for key model parameters that account for a linear increase with leaf N of V_max, J_max, T_p, and R_dmax (maximum rates of carboxylation, electron transport, triose phosphate export, and mitochondrial respiration), a saturation-type increase of φ (quantum yield of electron transport), and a non-linear decrease of θ and m (curvature of the light dependence of electron transport rate, scaling factor of the stomata model). The adaptation of photosynthetic characteristics to T_g was included into the model by linear relations that were observed between T_g and the activation energy ΔH_a of the temperature response characteristics of V_max, J_max, and T_p as well as of the nitrogen dependency of these characteristics. Based on an analysis of diurnal time courses of gas exchange rates it was found necessary including not only the relation between leaf water potential (Ψ) and stomatal conductance as used originally in LEAFC3, but additional effects on V_max and J_max. With the above-listed extensions, the model was capable to reproduce the observed plasticity and the recorded diurnal time courses of gas exchange rates fairly well. Thus, we conclude that the new model version can be used under a broad range of conditions, both for ecophysiological studies and as a submodel of crop growth models. The results presented here for barley will facilitate adapting photosynthesis models like LEAFC3-N to other C₃-species as well. The modelling of the effects of drought stress should be further elaborated in future based on more specific experiments.     

Trophic diversity in amphipods within a temperate eelgrass ecosystem as determined by gut contents and C and N isotope analysis

A conjoint analysis of gut contents and stable C and N isotopes was applied to determine the main food sources and feeding habits of dominant amphipods in an eelgrass bed (Zostera marina) in Gwangyang Bay, Korea. Gut content observations demonstrated that, while Gammaropsis japonicus and Jassa slatteryi are herbivorous, feeding on epiphytes and detritus, Pontogeneia rostrata and Monocorophium acherusicum are omnivorous, feeding on mesozooplankton fragments and detritus. Stable isotope data confirmed that epiphytes, detritus, and mesozooplankton fragments were major food sources for amphipods in the eelgrass bed. Isotopic mixing model calculations clearly showed an interspecific difference in diet composition. A high isotopic dissimilarity between amphipod taxa demonstrated interspecific trophic diversity, reflecting their herbivorous (G. japonicus and J. slatteryi) and omnivorous (P. rostrata and M. acherusicum) feeding habits and confirmed the detrivorous feeding habits of caprellids. Such trophic diversity at interspecific level of the amphipod species indicates that they use different food resources within their microhabitats and play species-specific functional roles as mediators in trophic pathways from producers to higher-level consumers of the eelgrass ecosystem. Finally, our findings suggest that information on the species-specific trophic ecology of amphipods is needed to better understand their potential role in the trophic dynamics and carbon flow of seagrass bed ecosystems.     

How does burrowing by the isopod <Emphasis Type="Italic">Limnoria agrostisa</Emphasis> (Crustacea: Limnoriidae) affect the leaf canopy of the southern Australian seagrass <Emphasis Type="Italic">Amphibolis griffithii</Emphasis>?

In south-western Australia, the isopod Limnoria agrostisa commonly burrows into leaf clusters and immature shoots of Amphibolis griffithii. The isopod also burrows into the sheath and rhizomes of Posidonia species. In A. griffithii, the isopod consumes new tissue within the sheath, damaging or destroying the meristem. This results in malformation of new leaves or destruction of whole leaf clusters with the potential to reduce the photosynthetic area of a shoot. The isopod has been found in all but one meadow of A. griffithii examined over 1,000 km of the Western Australian coastline. It was present throughout the year and showed little variation in abundance. Young were produced year round, but were more abundant in the summer months. Females, 3.5 mm in length or larger, produced 2–5 young that were brooded within the leaf cluster or base of an immature shoot. Within a meadow, 40–70% of shoots and 10–20% of leaf clusters were damaged by isopods. Seasonal trends were not consistent, but damage appeared to be higher in summer when isopod abundance was higher. Approximately 40% of clusters were destroyed by isopod damage. Isopods attack shoots of all ages, but damage was often located on apical clusters. There was no evidence that isopod damage initiated branching or leaf cluster formation. Estimations of clusters damaged or destroyed may be conservative, as only those clusters remaining on a shoot could be counted, and cluster loss could not be quantified. Examination of defoliated stems on upright shoots and horizontal rhizomes indicated that many were destroyed by isopods. The effect of L. agrostisa although substantial appears to be a feature of healthy seagrasses throughout southern Australia.     

Coupling biochemical and biophysical processes at the leaf level: an equilibrium photosynthesis model for leaves of C3 plants

The paper presents a generic computer model for estimating short-term steady-state fluxes of CO₂, water vapor, and heat from broad leaves and needle-leaved coniferous shoots of C₃ plant species. The model explicitly couples all major processes and feedbacks known to impact leaf biochemistry and biophysics including biochemical reactions, stomatal function, and leaf-boundary layer heat- and mass-transport mechanisms. The ability of the model to successfully predict measured photosynthesis and stomatal-conductance data as well as to simulate a variety of observed leaf responses is demonstrated. A model application investigating physiological and environmental regulation of leaf water-use efficiency (WUE) under steady-state conditions is discussed. Simulation results suggest that leaf physiology has a significant control over the environmental sensitivity of leaf WUE. The implementation of a highly efficient solution technique allows the model to be directly incorporated into plant-canopy and terrestrial ecosystem models.