Energy budget of the predatory marine gastropod Polinices duplicatus

Based on direct monitoring of feeding and growth under field conditions and frequent measurements of respiration and energy content in the laboratory, annual energy budgets were derived for Polinices duplicatus feeding on Mya arenaria at Barnstable Harbor, Massachusetts. The gross growth efficiency (P_g/C) of P. duplicatus was high (35%) and varied inversely with snail size. The proportion of ingested energy expended on respiration (R/C) was modest (44%) and varied directly with snail size and inversely with temperature. The energetic efficiencies of P. duplicatus agreed with the few results available for othe predatory molluscs and differed from most results available for herbivorous and detritivorous molluscs.     

Calcium and magnesium carbonate concentrations in different growth regions of gorgonians

Four of the most abundant gorgonian species from the southwestern Cape waters, Eunicella papillosa (Esper, 1797), E. alba (Esper, 1797), E. tricoronata Velimirov, 1971 and Lophogorgia flamea (Ellis and Solander, 1786) were analysed for Ca and Mg by atomic absorption spectroscopy (AAS) and ethylenediaminetetraacetate (EDTA) titration. The total mineral content in the peripheral tissues, excluding the axial skeleton, expressed as the sum of CaCO₃ and MgCO₃ of dry matter was between 65.5 and 83.5%. The mineral content varied in different growth regions and all specimens showed a higher degree of mineralization at the base than at the branch tips. The MgCO₃ concentration varied with genus and species and was between 9 and 11 mol %. The variation of the MgCO₃ concentration within different growth regions of the same species was small and generally did not exceed 0.8 mol %. From the branch to the stem, CaCO₃ and total mineral content was found to increase. The CaCO₃:MgCO₃ rations in different growth regions of all species indicated that the composition of the mesoskeleton with regard to the relative concentration of CaCO₃ and MgCO₃ is constant throughout the animal. Mineralogically, the mesoskeleton consists of high magnesian calcite as identified by X-ray diffraction. MgCO₃ concentrations determined by the peak shift method and by AAS were in fair agreement. The MgCO₃ data in gorgonian samples from the cold Atlantic Ocean and the warmer Indian Ocean show a linear relationship between water temperature and MgCO₃ concentration already demonstrated by Chave (1954). However, our data were consistently lower by 1 to 2% than expected.     

Suitability of estuarine nursery zones for juvenile weakfish (Cynoscion regalis): effects of temperature and salinity on feeding,growth and survival

Juvenile weakfish, Cynoscion regalis (Bloch and Schneider, 1801), exhibit significant spatial diffrences in growth rate and condition factor among estuarine nursery zones in Delaware Bay. The potential influence of temperature and salinity on the suitability of estuarine nursery areas for juvenile weakfish was investigated in laboratory experiments by measuring ad libitum feeding rate, growth rate and gross growth efficiency of juveniles collected in Delaware Bay in 1990 (40 to 50 mm standard length; 1.4 to 2.1 g) in 12 temperature/salinity treatments (temperatures: 20, 24, 28°C; salinities: 5, 12, 19, 26 ppt) representing conditions encountered in different estuarine zones during spring/summer. Feeding rates (FR) increased significantly with temperature at all salinities, ranging from 10 to 15% body wt d^-1 at 20°C to 33–39% body wt d^-1 at 28°C. Specific growth rates (SGR) ranged from 1.4 to 9.4% body wt d^-1 (0.3 to 1.5 mm d^-1) and gross growth efficiencies (K ₁) varied from 13.6 to 26.4% across temperature/salinity combinations. Based on nonlinear multiple regression models, predicted optimal temperatures for SGR and K ₁ were 29 and 27°C, respectively. Salinity effects on SGR and K ₁ were significant at 24 and 28°C where predicted optimal salinity was 20 ppt. At these warmer temperatures, SGR and K ₁ were significantly lower at 5 than at 19 ppt despite higher FR at 5 ppt. Therefore, maximum growth rate and growth efficiency occurred under conditions characteristic of mesohaline nurseries. This finding is consistent with spatial patterns of growth in Delaware Bay, implying that physicochemical gradients influence the value of particular estuarine zones as nurseries for juvenile weakfish by affecting the energetics of feeding and growth. Laboratory results indicate a seasonal shift in the location of physiologically optimal nurseries within estuaries. During late spring/early summer, warmer temperatures in oligohaline areas permit higher feeding rate and faster growth compared to mesohaline areas. By mid-late summer, spatial temperature gradients diminish and mesohaline areas provide more suitable physicochemical conditions for growth rate and growth efficiency whereas oligohaline areas become energetically stressful. Substantial mortality occurred at 5 ppt and 28°C, providing additional evidence that oligohaline conditions are stressful during late summer. Furthermore, juveniles provided a choice among salinities in laboratory trials preferred those salinities which promoted higher growth rates. The extensive use of oligohaline nurseries by juvenile weakfish despite the potential for reduced growth rate and growth efficiency suggests this estuarine zone may provide a substantial refuge from predation.     

Rearing Pandalus borealis larvae in the laboratory

Larvae of the northern shrimp Pandalus borealis (Krøyer) are pelagic. In the Estuary and Gulf of St. Lawrence, Canada, the early stages are found in the upper 25-m of the water column in spring and early summer and are expected to experience a range of water temperatures from as low as 0°C to as high at 6°C. Little is known of the impact of water temperature on metabolic requirements of northern shrimp larvae. In this study, routine respiration (VO₂), maximum respiration (electron transport system activity, ETSA) and metabolic scope for growth (MS, ETSA–VO₂) of northern shrimp larvae were measured as a function of temperature (3, 5 and 8°C), developmental stage (I–V at 3°C, I–VII at 5°C and 8°C) and growth rate in dry mass. After logarithmic transformation, all three metabolic variables were linearly related to dry mass. The increase in VO₂ with body mass was faster at 5°C than at 3 or 8°C, whereas with ETSA this increase was slower. As a result, MS increased more slowly with dry mass at 5°C than at 3 and 8°C. However, MS did not limit growth in this study, since it explained only 39% of the variability in growth. All three metabolic variables as well as growth varied together as a function of temperature and ontogeny. Q₁₀ of all three metabolic variables ranged from 1.6 and 2.2 for stages I–V larvae, except for VO₂ at stage I (3.9) and stage III (2.9).     

The influence of storage on antibacterial activity of sea water. I. Experiments with sea water stored at 18°C

Three sea-water samples of about 25 l each, stored at 18°C in the dark, were examined for antibacterial activity and bacterial content. Serratia marinorubra mainly was used as test bacterium. During storage for 197 to 454 days, bactericidal capacity varied considerably. In raw sea water, extreme log (N ₃–N ₀) values (N: number of colony-forming test bacteria/ml after 0 and 3 days of any test) were higher than-1 and lower than-4. Antibacterial activity of filtersterilized and autoclaved, stored, sea water likewise varied, but to a lesser degree. The number of colony-forming marine bacteria fluctuated about 3 orders of magnitude. Variations in bacterial content and in bactericidal action of stored sea water were positively correlated during the first months of storage; later, this correlation became negative. From these results it is concluded that variations in nutrient concentration — due to successive periods of growth and autolysis of marine bacteria — were the primary reason for the changes observed in antibacterial activity of stored sea-water samples.     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.     

Growth rate and carbon affinity ofUlva lactuca under controlled levels of carbon,pH and oxygen

We examined the growth rate (µ) ofUlva lactuca L. (collected from Roskilde Fjord, Denmark in 1987) at different levels of dissolved inorganic carbon (DIC), pH and oxygen in two culture facilities. Growth was faster in Facility A (µ _max ca 0.3 d^–1) than in B (µ _max ca 0.2 d^–1), probably because of more efficient stirring and higher light intensity. The growth-DIC response curve exhibited low half-saturation constant (K _1/2) values (0.35 mM DIC in A, 0.55 mM in B) and growth rates close toµ _max at natural seawater concentration of 2 mM DIC. Growth rate showed a low sensitivity to oxygen over a wide range of DIC and oxygen concentrations. Collectively, the results demonstrated an efficient mechanism for DIC use, unaffected by acclimatization to DIC concentrations between 0.2 and 3 mM. The growth rate decreased little between pH 7.5 and 9 at 2 mM DIC, but steeply above pH 9 approaching zero just above pH 10. The decline of growth at high pH may result from direct pH effects on cell pH, reduced HCO ₃ ^- availability and impaired operation of the carbon uptake process. The growth responses ofU. lactuca to DIC, pH and oxygen resembled those observed in previous short-term photosynthetic experiments. This similarity is probably due to the fast growth ofU. lactuca which means that photosynthetic products are rapidly converted into cell growth. Based on the culture experiments we argue that field plants ofU. lactuca not exposed to stagnant water and DIC depletion are likely to be limited in growth by environmental factors other than DIC (e.g. light and nutrients). Dense mats ofU. lactuca, however, may show reduced growth as a result of DIC depletion, high pH and self-shading.     

Energetics of Euphausia pacifica. I. Effects of body carbon and nitrogen and temperature on measured and predicted production

Laboratory production during the life span of Euphausia pacifica was measured directly (as the sum of growth, molting and reproduction) and indirectly (as assimilation minus metabolism and leakage) to test the hypothesis that weight-specific production is a constant for all sizes. Euphausiids were collected in Puget Sound, Washington State, USA, from September 1973 to March 1978. Equations were determined (in terms of carbon and nitrogen at 8° and 12° C) expressing the relationships between body weight and the daily rates of growth, molting, reproduction, ingestion and metabolism. The allometric equation (R=aW ^b ) best related body weight (W) to the rate (R) for growth, molting, ingestion, respiration and excretion for life stages from late larvae through adults. As predicted by the original above hypothesis, the weight-specific coefficient (b) was close to 1.0 for ingestion and excretion; in contrast, b was 0.62 for growth, and 0.77 to 0.85 for molting and respiration. The Q₁₀ s also varied: 3.5 for growth, 2.4 for molting, about 3.0 for ingestion, and 2.0 for respiration and excretion. Assimilation efficiencies, for all weights and at both temperatures, were 81.3% of carbon and 85.9% of nitrogen ingested. The relationships between rate and body weight of early larvae for growth and molting were linear, as was the relationship for reproduction in adults. Weight-specific production was higher by I to 2% at 12° than 8° C for all life stages, and was 2 to 4% for carbon and 2 to 6% for nitrogen in adults, but 13 to 17% for carbon and 14 to 15% for nitrogen in early furcilia larvae. The null hypothesis was rejected for production measured directly, but would have been accepted if only an indirect measurement of nitrogen production had been considered. Clearly, indirect measurement incorporates all errors of measurement and assumption and makes interpretation difficult.     

Effect of light and temperature on the population dynamics of two toxic bloom forming Cyanobacteria – Microcystis ichthyoblabe and Anabaena aphanizomenoides

The effect of light and temperature on the growth of Microcystis ichthyoblabe and Anabaena aphanizomenoides, isolated from the subtropical Oued Mellah lake, Morocco (33°30′N–07°20′W), were investigated in batch culture. Growth rates at 66 light–temperature combinations were determined and fitted with different mathematical models. The results show that the two Cyanobacteria grow at all light intensities and temperatures, except at 10 °C for A. aphanizomenoides, where the growth was strongly limited. The μ_max of M. ichthyoblabe increased with temperature from 0.56 d^?1 at 10 °C to 1.32 d^?1 at 35 °C. At all tested temperatures, a relative photoinhibition within the studied range of irradiance was observed and the photosensitivity was thermodependent. For Anabaena, the obtained μ_max ranged between 0.07 d^?1 at 10 °C and 1.46 d^?1 at 35 °C, and a weak photoinhibition was observed at 15 °C. The positive correlation between μ_max and I_opt (r²≥0.93) indicates a close interaction between light and temperature on the cyanobacteria growth. The results obtained in this work suggest that the growth of these two species is possible under low light and low temperature.     

Ingestion-independent rates of ammonium excretion by the copepod Calanus pacificus

The relationship between food ingested and NH ₊ ⁴ excretion rate was investigated for female Calanus pacificus collected in August, 1982, from the San Juan Archipelago, Washington State, USA. The copepods were preconditioned to 6 densities of the diatom Thalassiosira weissflogii (0 to 10⁴ cells ml^–1) for 30 h before the experiment. The experiment was conducted with nutrients added in excess to maintain equal rates of NH ₊ ⁴ uptake by the diatoms at all densities. Although ingestion rates of C. pacificus varied from 0 to over 20% of body N d^–1 at the different food levels, excretion was a constant 6.6 nM NH ₊ ⁴ copepod^–1 h^–1 or about 10% of body N d^–1. This ingestion-excretion relationship, which is consistent with previous respiration and fecundity studies, suggests that the ecological dominance of C. pacificus only under conditions of high food abundance may be due to a dramatic increase in its growth efficiency as ingestion increases above the level supporting a constant metabolic rate. The maintenance of a constant level of metabolism during relatively short periods of low food abundance may be advantageous if it allows the copepod to exploit more effectively short-term variability in its food resulting from environmental heterogeneity or vertical migration.Contribution No. 1360 from the School of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Measurement of calcium carbonate deposition in molluscs by controlled etching of radioactively labeled shells

Rates of calcium carbonate removal from shell pieces of Argopecten irradians (Say) and Mercenaria mercenaria (L.) were dependent on the type of etching fluid used and not shell origin. Etching was uniform over the entire shell surface, but surface morphology differed with etching fluids. Peak radioactivity was in early eluant fractions of shells etched immediately after radioactive labeling, and in later fractions when individuals were placed in isotope-free seawater after labeling. The etching technique can measure growth during the labeling period and subsequently by estimating the amount of calcium in fractions prior to the radioactive peak. Geometry of shell layers influenced the pattern of radioactivity seen in fractions. Peak location varied inside and outside the pallial line of individual M. mercenaria. A significant portion of the inorganic carbon used in shell formation by M. mercenaria was derived from metabolic CO₂.     

Influence of temperature on the growth rate of Skeletonema costatum in response to variations in daily light intensity

Daily light intensities (I _o) can vary 10-fold during the winter-spring and late-summer diatom blooms in New England, USA, coastal waters. Laboratory cultures and natural populations incubated in dialysis sacs were examined to determine the time course of growth rate in Skeletonema costatum (Greville) Cleve in response to variations in daily light intensity during two bloom periods in Narragansett Bay, Rhode Island, USA. Log-phase cultures of S. costatum require 2 d to attain maximum growth rates at 2°C following transfer to saturating intensities. At 20°C, only 1 d is required. As temperature increases, Detonula confervacea (Cleve) Gran, Thalassiosira nordenskiöldii Cleve and Ditylum brightwellii (West) Grunow also exhibit rapid increases in mean daily division rates (K) following transfer to saturating light intensities. Thalassiosira pseudonana Hustedt, however, did not alter the time required to achieve maximum K as temperature varied. Natural populations of S. costatum did not show a well-defined relationship between K and light. Throughout a winterspring bloom, K was limited by low temperatures and exhibited no clear response to variations in I _o. A change in K in response to variation in I _o may occur on a daily basis during the summer, when temperatures are near 20°C; this has yet to be verified for in situ populations.     

Use of high-performance liquid chromatography to investigate the production of paralytic shellfish toxins by Protogonyaulax spp. in culture

Procedures have been developed for the extraction and high-performance liquid chromatography (HPLC) analysis of paralytic shellfish poisoning (PSP) toxins from Protogonyaulax spp. grown in batch culture. Using these procedures, the toxin content of two isolates of P. tamarensis (NEPCC 183 and 255) and one isolate of P. catenella (NEPCC 355) were examined. Total toxin and individual toxin concentrations were measured for each isolate during the exponential and stationary phases of growth in batch culture. The total toxicity of each isolate as measured by HPLC analysis was found to agree with toxicity as determined by the standard mouse bioassay. Two of the isolates (255 and 355) were found to be toxic and the third (183) was non-toxic. The toxic isolates (255 and 355) both showed higher average total PSP toxin content during the exponential phase (35 and 23 fmol toxin cell^-1, respectively) than during the stationary phase (21 and 8 fmol toxin cell^-1, respectively). These cultures differed dramatically in their toxin composition. P. tamarensis (255) contained a large proportion of the N(21) sulfo toxins (B₁, B₂, C₁, C₂) while P. catenella (355) contained primarily Gonyautoxins 1 through 4. The percent composition of individual toxins was found to be constant throughout the growth cycle for both toxic isolates, even though the total toxin concentration varied. Our results suggest that PSP toxin profiles might be useful as chemotaxonomic indicators.     

Growth,production and ecological significance of Ophiura albida and O. ophiura (Echinodermata: Ophiuroidea) in the German Bight

Growth and production of the shallow-water ophiuroids Ophiura albida and O. ophiura were investigated at two stations in the German Bight from 1988 to 1991. Growth rings visible on the vertebral ossicles of the ophiuroid arms were interpreted as annual age markers. A correction for overgrown first rings allows for more exact estimations of growth and age. In both species growth could be described by Von Bertalanffy growth functions with the asymptotic disc diameter D =10.1 mm, K=0.229 and t _o=-0.192 in O. albida and D =27.7 mm, K=0.084 and t _o=0.042 in O. ophiura. Somatic production was calculated from mass specific growth rates. Annual production:biomass (P:B) ratios were estimated at 0.32 for O. albida and 0.43 for O. ophiura.AWI Publication Number: 618     

Energy allocation in a reef coral under varying resource availability

An organism’s pattern of resource allocation to reproduction and growth over time critically impacts on its lifetime reproductive success. During times of low resource availability, there are two fundamental, mutually exclusive strategies of energy investment: maintenance of somatic tissues to support survival and later reproduction or investment into an immediate reproductive event at the risk of subsequent death. Here, we examine energy allocation patterns in the coral Montipora digitata to determine whether energy investment during periods of resource shortage favours growth or reproduction in a sessile, modular marine species. We manipulated light regimes (two levels of shading) on plots within a shallow reef flat habitat (Orpheus Island, Great Barrier Reef, Australia) and quantified energy uptake (rates of net photosynthesis), energy investment into reproduction (E _R ), tissue growth per unit surface area (E _T ) and energy channelled into calcification (E _C ). With declining resource availability (i.e. reduced photosynthesis), relative energy investment shifted from high (~80%) allocation to tissue growth (E _R :E _T :E _C  = 11:81:8%) to an increasing proportion channelled into reproduction and skeletal growth (20:31:49%). At the lowest light regime, calcification was maintained but reproduction was halted and thus energy content per unit surface area of tissue declined, although no mortality was observed. The changing hierarchy in energy allocation among life functions with increasing resource limitation found here for an autotrophic coral, culminating in cessation of reproduction when limitations are severe, stands in contrast to observations from annual plants. However, the strategy may be optimal for maximising fitness components (growth, reproduction and survival) through time in marine modular animals.     

Growth of Chaetodon larvatus (Chaetodontidae: Pisces) in the southern Red Sea

Growth and age of Chaetodon larvatus were studied using growth bands in otoliths and length-frequency analyses. Otoliths of 180 C. larvatus were extracted and measured. Polished sections of sagittae revealed alternating opaque and translucent bands corresponding with a seasonal growth pattern. Both mass and size of the otoliths continue to grow steadily throughout life. Length-at-age data revealed very fast growth during the first year. Growth proceeded at a decreasing rate during the second and the third year; fishes older than 3 years did not grow noticeably. No difference in growth patterns between males and females could be detected. The growth parameters obtained for the whole population are: the asymptotic length (L _∞)=10.64 cm, growth constant (K)=1.14 year⁻¹ and the theoretical age at length zero (t ₀)=−0.30 year. The maximum age recorded was 14 years. Length frequency data collected at a recruitment site confirmed the fast growth of juveniles.     

Photoadaptation of photosynthesis in Gonyaulax polyedra

Gonyaulax polyedra Stein exhibited a combination of photoadaptive strategies of photosynthesis when only a single environmental variable, the light intensity during growth, was altered. Which of several biochemical/physiological adjustments to the light environment were employed depended on the level of growth irradiance. The photoadaptive strategies employed over any small range of light levels appeared to be those best suited for optimizing photosynthetic performance and not photosynthetic capacity. (Photosynthetic performance, P _i, is defined as the rate of photosynthesis occurring at the level of growth irradiance.) Among all photosynthetic parameters examined, only photosynthetic performance showed a consistent correspondence to growth rates of G. polyedra. Above 3500 to 4000 W cm^-2, where photosynthetic performance was equal to photosynthetic capacity, cells were not considered light-limited in either photosynthesis or growth. At these higher light levels, photosynthetic perfomance, cell volume, growth rates and respiration rates remained maximal; photosynthetic pigment content varied only slightly, while the photosynthetic capacity of the cells declined. At intermediate light levels (3000 to 1500 W cm^-2), photosynthesis, not growth, was light-limited, and photoadaptive strategies were induced which enhance absorption capabilities and energy transfer efficiencies of chlorophyll a to the reaction centers of G. polyedra. Photosynthetic capacity remained constant at about 280 mol O₂ cm^-3 h^-1, while photosynthetic performance ranged from 100 to 130 mol O₂ cm^-3 h^-1. Major increases in photosynthetic pigments, especially peridinin-chlorophyll a-proteins and an unidentified chlorophyll c component, accompanied photoadaptation to low irradiances. Maximal growth rates of 0.3 divisions day^-1 were maintained, as were respiration rates of about-80 mol O₂ cm^-3 h^-1 and cell volumes of about 5.4×10^-8 cm^-3 cell^-1. Below about 1250 W cm^-2, photosynthesis in G. polyedra was so light-limited that photosynthetic performance was unable to support maximal growth rates. Under these conditions, G. polyedra displayed photostress responses rather than photoadaptive strategies. Photostress was manifested as reduced cell volumes, slower growth, and drastic reductions in pigmentation, photosynthetic capacity, and rates of dark respiration.     

Moderate seawater acidification does not elicit long-term metabolic depression in the blue mussel Mytilus edulis

Marine organisms are exposed to increasingly acidic oceans, as a result of equilibration of surface ocean water with rising atmospheric CO₂ concentrations. In this study, we examined the physiological response of Mytilus edulis from the Baltic Sea, grown for 2 months at 4 seawater pCO₂ levels (39, 113, 243 and 405 Pa/385, 1,120, 2,400 and 4,000 μatm). Shell and somatic growth, calcification, oxygen consumption and \textNH₄ ⁺ {\text{NH}}_{4}^{ + } excretion rates were measured in order to test the hypothesis whether exposure to elevated seawater pCO₂ is causally related to metabolic depression. During the experimental period, mussel shell mass and shell-free dry mass (SFDM) increased at least by a factor of two and three, respectively. However, shell length and shell mass growth decreased linearly with increasing pCO₂ by 6–20 and 10–34%, while SFDM growth was not significantly affected by hypercapnia. We observed a parabolic change in routine metabolic rates with increasing pCO₂ and the highest rates (+60%) at 243 Pa. \textNH₄ ⁺ {\text{NH}}_{4}^{ + } excretion rose linearly with increasing pCO₂. Decreased O:N ratios at the highest seawater pCO₂ indicate enhanced protein metabolism which may contribute to intracellular pH regulation. We suggest that reduced shell growth under severe acidification is not caused by (global) metabolic depression but is potentially due to synergistic effects of increased cellular energy demand and nitrogen loss.     

PIXGRO: A model for simulating the ecosystem CO2 exchange and growth of spring barley

A model, PIXGRO, developed by coupling a canopy flux sub-model (PROXEL_NEE; PROcess-based piXEL Net Ecosystem CO₂ Exchange) to a vegetation structure submodel (CGRO), for simulating both net ecosystem CO₂ exchange (NEE) and growth of spring barley is described. PIXGRO is an extension of the stand-level CO₂ and H₂O-flux model PROXEL_NEE, that simulates the NEE on a process basis, but goes further to include the dry matter production, partitioning, and crop development for spring barley. Dry matter partitioned to the leaf was converted to leaf area index (LAI) using relationships for the specific leaf area (SLA). The canopy flux component, PROXEL_NEE was calibrated using information from the literature on C3 plants and was tested using CO₂ flux data from an eddy-covariance (EC) method in Finland with long-term observations. The growth component (CGRO) was calibrated using data from the literature on spring barley as well as data from the Finland site. It was then validated against field data from two sites in Germany and partly via the use of MODIS remotely sensed LAI from the Finland site.Both the diurnal and the seasonal patterns of gross CO₂ uptake were very well simulated (R² = 0.92). A slight seasonal bias may be attributed to leaf ageing. Crop growth was also well simulated; simulated dry matter agreed with field observed data from Germany (R² = 0.90). For LAI, the agreement between the simulated and observed was good (R² = 0.80), giving an indication that functions describing the conversion of fixed CO₂ to dry matter and the subsequent partitioning leaf dry matter and LAI simulation were robust and provided reliable estimates.The MODIS LAI at a resolution of 1000 m agreed poorly (R² = 0.45) with the PIXGRO simulated LAI and the observed LAI at the Finland site in 2001. We attributed this to the coarse resolution of the image and/or the small size of the barley field (about 17 ha or 0.25 km²) at the Finland site. By deriving a regression relation between the observed LAI and NDVI from a higher resolution MODIS (500 m resolution), the MODIS-recalculated LAI agreed better with the PIXGRO-simulated LAI (R² = 0.86).PIXGRO provides a prototype model bridging the disciplines of plant physiology, crop modeling and remote sensing, for use in a spatial context in evaluating carbon balances and plant growth at stand level, landscape, regional, and with some care, continental scales. Since almost 50% of the European land surface is covered by crops, such a model is needed for the dynamic estimation of LAI and NEE of croplands.