Seasonal nitrogen speciation in temperate seagrass Posidonia oceanica (L.) Delile

To better understand some basic aspects of the nitrogen economy in Posidonia oceanica and, specifically, the seasonality of the processes of storage, translocation and assimilation, we examined nitrogen speciation into soluble compounds, both inorganic (nitrates, nitrites and ammonium) and organic (free amino acids, FAA, and total soluble protein, TSP), and the nitrogen assimilation potential (through the glutamine synthetase activity measurement) in the leaves, rhizomes and roots of P. oceanica over a 1-year cycle. Only a limited amount of inorganic nitrogen was found, accounting for less than 3.3% of the total nitrogen content, and it was mostly in the form of ammonium. Nitrate and nitrite concentrations were very low, always below 7.2 μmol g⁻¹ dw in annual average. Among the organic soluble fractions, FAAs were the most abundant, accounting for up to 50% of N pools. Rhizomes were the organs in which FAA concentrations reached their maximum value. The leaves showed higher nitrogen assimilation potential than the roots and this assimilation potential was highest during and after the period of maximum leaf growth, probably corresponding to the assimilation of both new and recycled nitrogen. Our results suggest that 5% of the total nitrogen assimilation occurs in roots and 79% in leaves on an annual average. In addition, rhizomes contributed to the total shoot nitrogen assimilation by 32-54% between autumn and spring. Rhizomes appear as key organs in the nitrogen economy of the plant, not only as a major site for nitrogen assimilation but also as an organ for nitrogen storage. This storage, mostly in the form of FAA, occurs during periods of high availability and low demand (winter). This stored nitrogen can supply up to 33% of plant demands during the moment of maximum leaf growth (i.e. late spring).     

Strategies of nitrogen cycling of Phragmites australis at two sites differing in nutrient availability

Below-ground biomass and nitrogen content were determined at two genetically homogeneous Phragmites stands differing in morphology, in productivity, and in nutrient supply. Comparable ratios between above-ground standing crop and rhizome biomass were found at both sites, whereas the root biomass/above standing crop ratio was significantly higher at the nutrient poor site. Investigations on the dynamics of nitrogen content revealed distinct differences in nitrogen translocation to the rhizomes between the investigated clones indicating two ecophysiological strategies in storage behaviour. These two strategies could be attached to the “assimilation type” and to the “translocation type”, respectively. A modified definition of both types is presented.     

Nitrogen metabolism of Douglas fir and Scots pine as affected by optimal nutrition and water supply under conditions of relatively high atmospheric nitrogen deposition

Nitrogen metabolism of the needles of 40-year-old Douglas fir and Scots pine trees, growing in two forest stands on cation-poor and acidic sandy soil with a relatively high atmospheric nitrogen deposition was studied. The composition of the free amino acid (FAA) pool, the concentrations of total nitrogen and soluble protein and the activities of glutamine synthetase (GS) and glutamate dehydrogenase (GDH) were determined in the needles. An excessive nitrogen supply by a high atmospheric nitrogen deposition in both forest stands was indicated by the high concentrations of total nitrogen and the amino acids arginine, glutamic acid, glutamine and aspartic acid in control trees. In addition the effect of optimal nutrition and water supply (fertigation) on the needle nitrogen metabolism was evaluated. The total concentration of the FAA pool in needles of both tree species was lower in the fertigated than in the non-fertigated (control) trees, except for 1-year-old needles of Scots pine, in which the concentration after fertigation did not differ from the control. The lower total FAA concentration in the fertigated trees could be attributed to arginine, the concentration of which was on average 60% lower than in the control. Neither the concentration of soluble protein nor the activity of GS were influenced by fertigation. The activity of GDH in fertigated trees only differed significantly from the control in October. Scots pine needles had higher concentrations of protein (50%) and higher activities of GS (44%) and GDH (25%) than Douglas fir needles. Possible explanations for the lower vitality of Douglas fir compared to Scots pine are given.     

Biomass Allocation in Cyperus papyrus in a Tropical Wetland,Lake Naivasha,Kenya1

The nutrient status of Lake Naivasha, a freshwater lake in southeastern Kenya, has been rising since at least 1982. A potential effect of increases in nutrient supply to the lake's floating papyrus is increasing of the plants’ investment in above–water material and reduction of the amount of energy invested in uptake and storage. Biomass and its allocation between culms, panicles, roots, and rhizomes was measured in 17 sites around the 150–km² lake. Although above–water biomass was greatest in sites closest to the lake's major nutrient inflow, the River Malewa, there was little evidence of corresponding decreases in the biomass of uptake and storage tissues. In August 1995, the average ± SD biomass of papyrus in the lake was 11,540 ± 3020 g/m², with the papyrus containing about 4500 ± 1900 g total carbon/m² and about 100 ± 70 g total nitrogen/m². Plant nitrogen contents did not vary with distance from the main external nutrient supply. Together with low nitrogen concentrations in the plants (0.60 ± 0.26 in culms and 0.99 ± 0.50% in rhizomes), very high carbon to nitrogen ratios (49 ± 20:1) and nitrogen fixation in the rhizosphere explaining only about half of the plants’ nitrogen, papyrus is a likely net sink for nitrogen supplied from the lake's increasingly cultivated watershed. Despite this role, clearance of papyrus in favor of agriculture partly explains the reductions in the area of papyrus within the lake basin from 48 km² in the late 1960s to 14 km² in 1995.     

Effects of nitrogen addition on nitrogen metabolism and carbon reserves in the temperate seagrass Posidonia oceanica

The effect of repeated N additions on a dense, shallow meadow of Posidonia oceanica (L.) Delile in the NW Mediterranean was studied over a year. N was added biweekly both to the sediment and to the water column as ammonium and nitrate. The most obvious result of these additions was an overall increase in N content (% DW) in all tissues of fertilized plants; this increase was maximum in rhizomes, with values of 5% N reached, which confirmed the storage capacities of these organs.Fertilization affected the different N fractions in distinct ways. The free amino acid (FAA) concentration increased the most, particularly in rhizomes and roots, suggesting the function of these compounds for N storage and, probably, translocation. The non-soluble N fraction also increased greatly. The total soluble protein (TSP) and the inorganic N forms concentrations were less sensitive to fertilization, and only increased moderately in a few cases. N assimilation, assessed through in vivo glutamine synthetase (GS) activity, was maximum in leaves after the peak of growth, which coincided with the lowest N values in both control and fertilized plants. Thus assimilation was probably greatest at the period of highest N deficiency. Growth rates did not respond to N enrichment. Another clear effect of N addition was to decrease carbon reserves. In effect, the concentration of total non-structural carbohydrate (TNC) greatly decreased in rhizomes of fertilized plants, coinciding with the increase in FAA. We conclude that increased nitrogen availability can affect plant survival through the decrease in their carbon reserves, crucial for P. oceanica overwintering. This interaction between N and C metabolism helps to explain changes in benthic vegetation after steadily increasing coastal water eutrophication.     

SEASONAL NUTRIENT DYNAMICS,NUTRIENT RESORPTION,AND MYCORRHIZAL INFECTION INTENSITY OF TWO PERENNIAL FOREST HERBS

Patterns of mycorrhizal infection, seasonal foliar nutrient concentrations, nutrient allocation to shoots and rhizomes, and nutrient resorption were measured in relation to soil nutrient availability in two species of perennial forest herbs, Geranium maculatum L. and Polygonatum pubescens Pursh., in four forest stands in southern and central Ohio. The percentage of plants with V-A mycorrhizae and the proportion of root length colonized by VAM structures increased with decreasing nutrient availability in both species. Foliar N and P concentrations in plants from lower fertility sites were as high, or higher, than those in plants from higher fertility sites; as a result, tissue nutrient enrichment ratios (foliar concentration/soil available concentration) increased with decreasing fertility. Proportional resorption of N and P generally decreased with decreasing nutrient availability, a pattern inconsistent with those exhibited by woody plants in these forest stands. We hypothesize that the inverse relationship between nutrient uptake efficiency (via mycorrhizae) and nutrient use efficiency (resorption) exhibited by these forest understory herbs, but not by trees or herbs from high-light environments, may be related to low-light limitation of energy reserves in the forest understory.     

Growth and allocation of the arctic sedges Eriohorum angustifolium and E. vaginatum: effects of variable soil oxygen and nutrient availability

In arctic tundra soil, oxygen depletion associated with soil flooding may control plant growth either directly through anoxia or indirectly through effects on nutrient availability. This study was designed to evaluate whether plant growth and physiology of two arctic sedge species are more strongly controlled by the direct or indirect effects of decreased soil aeration. Eriophorum angustifolium and E. vaginatum, which originate from flooded and well-drained habitats, respectively, were grown in an in situ transplant garden at two levels of soil oxygen, nitrogen, and phosphorus availability over two growing seasons. In both species, N addition had a stronger effect on growth and biomass allocation than P addition or soil oxygen depletion. Net photosynthesis and carbohydrate concentrations were relatively insensitive to changes in these factors. Biomass reallocated from shoots to below-ground parts in response to limited N supply was equally divided between roots (nutrient acquisition) and perennating rhizomes (storage tissue formation) in E. angustifolium. E. Vaginatum only increased its allocation to rhizomes. In the flood-tolerant E. angustifolium, growth was improved by soil anoxia and biomass allocation among plant parts was not significantly affected. Contrary to our initial hypothesis, whole-plant growth in E. vaginatum improved in flooded soils; however, it only did so when N availability was high. Under low N availability growth in flooded soils was reduced by 20% compared to growth in the aerobic environment. Reduced biomass allocation to rhizomes and thus to storage potential under anaerobic conditions may reduce long-term survival of E. vaginatum in flooded habitats.     

Rhizome age structure of three populations ofPhragmites australis (Cav.) Trin. ex Steud.: Biomass and mineral nutrient concentrations

The rhizome age structure for populations ofPhragmites australis (Cav.) Trin. exSteud. was determined at three sites of the T?eboň Biosphere Reserve (Czech Republic). These sites were classified according to plant species composition and phosphorus availability as oligotrophic (Branná), mesotrophic (Ro?mberk East) and hypertrophic (Ro?mberk West).P.australis stands at these sites were expanding, stabilized and retreating, respectively. Rhizomes were sampled within the terrestrial parts of the reed stands (at a water depth of about 10 cm). The rhizomes were dated according to the position in the branching hierarchy and to supplementary morphological criteria. The analysis of the branching pattern revealed that at the sites studied, the rhizomes typically branch only once a year. The longest life span of rhizomes (six years) was found for the stable reed population at Ro?mberk East. The reduced rhizome life span at Ro?mberk West (four to five years), accompanied by a reduced branching frequency of horizontal rhizomes, was ascribed to a greater incidence of anaerobic conditions associated with the permanent flooding of the hypertrophic organic soil. The maximum life span of the Branná population (four years) probably corresponds to the total age of the reed population. The concentrations of nitrogen, phosphorus, and potassium in the tissue decreased with rhizome age while those of calcium and magnesium increased. The effect of rhizome age on mineral-nutrient concentrations was more pronounced than the effect of site on these concentrations. The mineral nutrient concentrations were not closely associated with reed performance.     

Soil Nitrogen Conditions Approach Preinvasion Levels following Restoration of Nitrogen-Fixing Black Locust (Robinia pseudoacacia) Stands in a Pine–Oak Ecosystem

Restoring native plant communities on sites formerly occupied by invasive nitrogen‐fixing species poses unique problems due to elevated soil nitrogen availability. Mitigation practices that reduce available nitrogen may ameliorate this problem. We evaluated the effects of tree removal followed by soil preparation or mulching on native plant growth and soil nitrogen transformations in a pine–oak system formerly occupied by exotic nitrogen‐fixing Black locust (Robinia pseudoacacia) trees. Greenhouse growth experiments with native grasses, Andropogon gerardii and Sorghastrum nutans, showed elevated relative growth rates in soils from Black locust compared with pine–oak stands. Field soil nutrient concentrations and rates of net nitrification and total net N‐mineralization were compared 2 and 4 years since Black locust removal and in control sites. Although soil nitrogen concentrations and total net N‐mineralization rates in the restored sites were reduced to levels that were similar to paired pine–oak stands after only 2 years, net nitrification rates remained 3–34 times higher in the restored sites. Other nutrient ion concentrations (Ca, Mg) and organic matter content were reduced, whereas phosphorus levels remained elevated in restored sites. Thus, 2–4 years following Black locust tree removal and soil horizon mixing achieved through site preparation, the concentrations of many soil nutrients returned to preinvasion levels. However, net nitrification rates remained elevated; cover cropping or carbon addition during restoration of sites invaded by nitrogen fixers could increase nitrogen immobilization and/or reduce nitrate availability, making sites more amenable to native plant establishment.     

Regrowth dynamics of <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis> after defoliation as affected by nitrogen availability

Young plants of a rhizomatous grass Calamagrostis epigejos (L.) Roth were grown from seed in nutrient solutions containing nitrogen in concentrations 0.1, 1.0, and 10 mM. After six weeks of cultivation the plants were defoliated and changes in growth parameters and in content of storage compounds were measured in the course of regrowth under highly reduced nitrogen availability. Plants grown at higher nitrogen supply before defoliation had higher amount of all types of nitrogen storage compounds (nitrates, free amino acids, soluble proteins), which was beneficial for their regrowth rate, in spite of lower content of storage saccharides. Amino acids and soluble proteins from roots and stubble bases were the most important sources of storage compounds for regrowth of the shoot. Faster growth of plants with higher N content was mediated by greater leaf area expansion and greater number of leaves. In plants with lower contents of N compounds number of green leaves decreased after defoliation significantly and senescing leaves presumably served as N source for other growing organs. Results suggest that internal N reserves can support regrowth of plants after defoliation even under fluctuating external N availability. Faster regrowth of C. epigejos with more reserves was mediated mainly by changes in plant morphogenesis.     

Mineral allocation to reproduction in Sorhum bicolor and Sorghum halepense in relation to parental nutrient supply

Summary This experiment investigated the effect of parental nutrient shortage on the allocation of five nutrients to seeds and rhizomes in Sorghum halepense, a perennial, noxious weed, and to seeds in Sorghum bicolor, an annual, cultivated species. Plants from both species were grown from seeds and supplied with fertilizer at three concentrations. The allocation of biomass and nutrients (N, P, K, Ca and Mg) to reproductive and vegetative parts was determined. Relative biomass allocation to reproduction (either sexual or vegetative) remained constant in S. halepense in spite of large differences in total plant weight. In S. bicolor, however, biomass allocation to sexual reproductive structures decreased significantly with decreasing nutrient supply. Individual seed weight was not modified by parental nutrient supply in S. halepense, but it increased with decreasing nutrient availability in S. bicolor. Important differences in mineral allocation to seeds were found between the two species. While S. bicolor seeds were largely buffered from the differences in parental nutrient status, concentration of nutrients in S. halepense seeds decreased significantly with decreasing supply for all the nutrients analyzed except Ca. However, mineral nutrient concentration in S. halepense rhizomes remained remarkably constant despite differences in the external supply, evincing the priority given to vegetative reproduction at the expense of sexual reproduction. Overall, the pattern of nutrient allocation in S. bicolor seeds under different nutrient supply resembled the pattern observed in S. halepense rhizomes, but it had little resemblance to the pattern of nutrient allocation in S. halepense seeds. The results are discussed in terms of differences and similarities in the reproductive strategy of these two species.     

根茎型植物扁秆荆三棱对光照强度和养分水平的生长响应及资源分配策略

自然界中光照和养分因子常存在时空变化,对植物造成选择压力。克隆植物可通过克隆生长和生物量分配的可塑性来适应环境变化。尽管一些研究关注了克隆植物对光照和养分因子的生长响应,但尚未深入全面了解克隆植物对光照和养分资源投资的分配策略。以根茎型草本克隆植物扁秆荆三棱(Bolboschoenus planiculmis)为研究对象,在温室实验中,将其独立分株种植于由2种光照强度(光照和遮阴)和4种养分水平(对照、低养分、中养分和高养分)交叉组成的8种处理组合中,研究了光照和养分对其生长繁殖及资源贮存策略的影响。结果表明,扁秆荆三棱的生长、无性繁殖及资源贮存性状均受到光照强度的显著影响,在遮阴条件下各生长繁殖性状指标被抑制。且构件的数目、长度等特征对养分差异的可塑性响应先于其生物量积累特征。在光照条件下,高养分处理的总生物量、叶片数、总根茎分株数、长根茎分株数、总根茎长、芽长度、芽数量等指标大于其他养分处理,而在遮阴条件下,其在不同养分处理间无显著差异,表明光照条件可影响养分对扁秆荆三棱可塑性的作用,且高营养水平不能补偿由于光照不足而导致的生长能力下降。光照强度显著影响了总根茎、总球茎及大、中、小球茎的生物量分配,遮阴条件下,总生物量减少了对地下部分根茎和球茎的分配,并将有限的生物量优先分配给小球茎。总根茎的生物量分配未对养分发生可塑性反应,而随着养分增加,总球茎分配下降,说明在养分受限的环境中球茎的贮存功能可缓冲资源缺乏对植物生长的影响。在相同条件下,根茎生物量对长根茎的分配显著大于短根茎,以保持较高的繁殖能力;而总球茎对有分株球茎的生物量分配小于无分株球茎,表明扁秆荆三棱总球茎对贮存功能的分配优先于繁殖功能。研究为进一步理解根茎型克隆植物对光强及基质养分环境变化的生态适应提供了依据。     

Plant biomass and nutrient levels of a tropical macrophyte (Cyperus papyrus L.) receiving domestic wastewater

The aboveground biomass and nutrient content ofCyperus papyrus were determined in a small tropical swamp receiving domestic wastewater. It was found that the biomass (4,955 g.m^–2 dry weight) was the highest ever reported for papyrus. The levels of both nitrogen and phosphorus in the plant organs were very high. The nitrogen concentrations of the various plant organs were 4.8% roots, 8.4% rhizomes, 4.5% scales, 4.8% culms, and 6.2% umbels on dry weight basis. As to phosphorus the concentrations were 0.09% roots, 0.11% rhizomes, 0.09% scales, 0.10% culms, and 0.13% umbels. The high biomass and nutrient contents of the plants may have been caused by the high nutrient levels in the surrounding water. Comparison of the nitrogen to phosphorus ratios in the plants to those in the surrounding water showed that the plants stored very high amounts of nitrogen.     

Nitrogen uptake by rhizomes of the clonal sedge Carex bigelowii: a previously overlooked nutritional benefit of rhizomatous growth

Studies on the rhizomatous clonal sedge Carex bigelowii , at a fellfield site in subarctic Swedish Lapland examined the ecological potential of rhizomes as nitrogen uptake systems. Direct application of a solution of ¹⁵NH₄¹⁵NO₃ to the rhizomes of C. bigelowii significantly enriched the ¹⁵N content of the plant tissue. The pattern of enrichment indicated movement of labelled nitrogen into rhizomes and adjoining tissue (including roots and shoots), showing both uptake and translocation via the rhizome system. There was a gradient of decreasing ¹⁵N enrichment with increasing distance from the point of labelling, and estimation of the total mass of nitrogen taken up via labelled rhizomes showed low levels and rates of uptake. Quantification of the size of rhizome and root systems of C. bigelowii at the study site indicates that 80% of the living biomass may be below ground, and that the surface area of the rhizome system of C. bigelowii is approx. two-thirds that of the roots. The rhizome system of C. bigelowii can therefore act as a route for nitrogen uptake, with the potential to exploit almost as great a volume of soil as the root system. This mechanism of nitrogen uptake may play an important role in the Arctic, where many species have a clonal, rhizomatous growth form. In addition, plant growth in many arctic ecosystems is limited by low soil nutrient availability, a result of low temperatures (leading to slow soil decomposition rates) and patchy resource distribution. Nitrogen uptake via rhizomes may provide plants with the capacity to take advantage of transient nutrient supplies, and may partly compensate for the cost of developing and maintaining persistent rhizome networks in ecosystems where nutrient resources are in short supply.     

Does ammonium affect accumulation of starch in rhizomes ofPhragmites australis (Cav.) Trin. ex Steud.?

Reed rhizomes from four habitats differing in soil nitrogen availability were collected and analysed for carbohydrate content during one year. Rhizomes collected from sediments where nitrate was the prevailing form of available nitrogen accumulated predominantly starch. On the other hand, rhizomes buried in sediments rich in ammonium contained predominantly sucrose. These observations together with the fact that ammonium stimulates the activity of enzymes involved in degradation of starch suggest that ammonium affects starch storage by inducing conversion of starch into sucrose. During spring when sprouting takes place, the rates of sucrose or starch mobilization differed remarkably, the sucrose in rhizomes buried in ammonium-rich sediments being depleted much faster than the starch in rhizomes at ammonium-poor habitats. The consequent reduction of total carbohydrate content in rhizomes at ammonium-rich habitats places the plants in a vulnerable situation. Eutrophication and reed decline are postulated to be events causally related via toxicity of ammonium, of which considerable amounts are microbiologically released from decaying organic matter in reed beds under excessive nutrient supply.     

南亚热带格木、马尾松幼龄人工纯林及其混交林生态系统碳氮储量

研究比较了南亚热带6年生格木(Erythrophleum fordii)、马尾松(Pinus massoniana)幼龄人工纯林及马尾松与格木混交林生态系统碳氮储量及其分配特征。结果表明,生态系统总碳储量依次为马尾松-格木混交林(137.75 t/hm2)格木纯林(134.07 t/hm2)马尾松纯林(131.10 t/hm2),总氮储量则为格木纯林(10.19 t/hm2)马尾松-格木混交林(8.68 t/hm2)马尾松纯林(7.01 t/hm2)。3种人工林生态系统碳氮库空间分布基本一致,绝大部分储存于0—100 cm土壤层,平均占生态系统总储量的81.49%和96.91%,其次为乔木层(分别占17.52%和2.69%),林下植被和凋落物层所占比例最小。林地土壤碳主要集中于表土层,其中0—30 cm土层平均碳储量为52.52 t/hm2,占土壤总碳储量(0—100 cm)的47.99%,土壤氮的分布则无明显规律。相比于纯林,与固氮树种混交的营林方式表现出更大的碳储存能力。3种幼龄人工林生态系统较低的地上与地下部分碳氮分配比,表明其仍具有较强的碳氮固持潜力。     

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Dissolved organic carbon (DOC) and nitrogen (DON) derived from aquatic and terrestrial vascular plants provide a major energy and nutrient source for freshwater and coastal marine biota. The bioavailability of this material may to a large extent depend on plant species. In this study, we have compared the bioavailability of DOC and DON sampled in two distinct stands of Typha domingensisand Eleocharis mutatain a coastal tropical lake and in the adjacent ocean in the state of Rio de Janeiro, Brazil. Bioavailability of organic matter was assessed by regrowth bioassays using natural bacterial inocula. Nutrients were added to achieve carbon or nitrogen limitation. At all sampling sites, DON comprised over 95% of the total bioavailable nitrogen, suggesting its dominant role as a nitrogen source. The bioavailability of lacustrine DON (22% in the Typhastand and 34% in the Eleocharisstand) exceeded the bioavailability of DOC (8 and 10%, respectively) and exhibited a larger difference between the stands. ³H-leucine incorporation studies showed that lake bacterioplankton had a well balanced supply of C, N and P. Therefore, an accumulation of labile DON due to an excess nitrogen supply is not probable. We propose that a substantial part of the lake DON was newly formed within the macrophyte stands, while DOC was predominantly of terrestrial origin and more diagenetically changed.     

Diel vertical migration thresholds of Karenia brevis (Dinophyceae)

Light and nutrient availability change throughout dinoflagellate diel vertical migration (DVM) and/or with sub-population location in the water column along the west Florida shelf. Typically, the vertical depth of the shelf is greater than the distance a sub-population can vertically migrate during a diel cycle, limiting the ability of a sub-population to photosynthetically fix carbon toward the surface and access nutrients sub-surface. This project investigated changes of Karenia brevis (C.C. Davis) G. Hansen et Moestrup intracellular carbon, nitrogen, internal nitrate (iNO₃), free amino acid (FAA), and total lipid concentrations in high-light, nitrate-replete (960 μmol quanta m⁻² s⁻¹, 80 μM NO₃), and high-light, nitrate-reduced (960 μmol quanta m⁻² s⁻¹, <5 μM NO₃) mesocosms. The nitrate-reduced mesocosm had a slowed cell division rate when compared to the nitrate-replete mesocosm. Minimum intracellular carbon, nitrogen, iNO₃, FAA, and total lipid concentrations during the largest surface sub-population aggregations led to the conclusion that daughter cells resulting from cell division received unequal shares of the parental resources and that this inequality influenced migration behavior. Nutrient reduced daughter cells were more strongly influenced by light and phototaxis for carbon production than their replete same cell division sister cells during vertical migration thus rapidly increasing the fulfillment of constituents through photosynthesis. Vertical migration was consistent with an optimization scheme based on threshold limits through utilization or formation of photosynthate. We propose a simplified conceptual model describing how K. brevis is transported along the benthos of the west Florida shelf from off-shore to on-shore. Dynamic carbon thresholds are also suggested for future DVM modeling efforts on K. brevis populations transported between nitrogen replete and nitrogen reduced environmental conditions.     

Nitrogen translocation via rhizome systems in monoclonal stands ofReynoutria japonica in an oligotrophic desert on Mt Fuji: Field experiments

Mechanisms which enableReynoutria japonica, a dominant pioneer herb, to be successful in maintaining large stands in an oligotrophic volcanic desert on Mt Fuji were investigated with special reference to its nitrogen acquisition.Reynoutria japonica forms circular stands, each of which comprises only one genet. As a stand develops outwards, the number of aerial shoots per unit area decreases in the center. Shoots grow vigorously in the peripheral area where the available nitrogen from soil and precipitation (about 2.4 g m⁻² year⁻²) was much less than total nitrogen in the shoots (6.1–9.1 g m⁻²). Leaf nitrogen content per unit mass was also greater in the leaves of the peripheral shoots. When rhizomes extending radially from the center to the periphery were severed, the dry mass of shoots in the periphery diminished by 75% on a ground area basis. In the periphery, leaf nitrogen content also reduced significantly and no flowers were produced. When fertilizer was applied to the peripheral shoots with severed rhizomes, neither growth, survival nor flower production of the shoots was significantly smaller than the control levels. In these shoots, it is also found that the nitrogen content in the youngest leaves decreased for about 1 month and then increased to above that in the control leaves. These results suggest that (i) nitrogen accumulated in the central part is translocated to peripheral shoots via rhizomes, and that long-distance translocation enables the stands to develop outwards, and (ii) aerial shoots in the periphery utilize the nitrogen translocated by rhizomes in the beginning of the growth season, whereas once the shoots have established, they begin to take up nitrogen with their own roots. Since the peripheral shoots are in sunnier environments than the shoots inside the stand, the acropetal nitrogen translocation via rhizomes will raise the production efficiency of a whole stand.     

Ribulose bisphosphate carboxylase, protein and nitrogen in Scots pine seedlings cultivated at different nutrient levels

Seedlings of Scots pine (Pinus sylvestris L.) of a northern provenance were cultivated in nutrient solution for 10 weeks in a climate chamber. The nutrient solution (renewed by solution exchange) contained 2.5, 10 or 50 mg N I^?1. All other essential elements were added in optimal proportion to the nitrogen. Seedlings cultivated at 10 and 50 mg N I^?1 were similar with respect to all characteristics studied. Seedlings cultivated at 2.5 mg N I^?1 showed a lower growth rate, especially for the shoot, and an altered morphology, with high root:shoot ratios and long, slender roots. The nitrogen concentrations in shoot and needles as well as in whole seedlings were not significantly affected by the nitrogen supply, while the nitrogen concentrations in the roots were somewhat lower at 2.5 mg N I^?1. Ribulose bisphosphate carboxylase (EC 4.1.1.39) activity and the concentrations of carboxylase, total and soluble protein and of chlorophyll in the needles were consistently much lower for seedlings cultivated at 2.5 mg N I^?1, than for seedlings grown at higher nutrient levels. A close correlation was observed between activity and concentration of the carboxylase (r=0.95). Carboxylase activity and protein were more sensitive to a low nutrient supply than was chlorophyll. The data show how activity and concentration of ribulose bisphosphate carboxylase and the concentrations of soluble and total protein and of chlorophyll in needles of pine seedlings can be negatively affected by the nutrient supply, also when the nitrogen concentrations in the needles are close to those observed at optimal nutrient supply. It is suggested that pine seedlings store assimilated non-protein nitrogen in the needles when protein synthesis is under restraint. The nitrogen concentration in needles and seedlings could not be used as a measure of the physiological state of the seedlings.