Nutrient addition does not increase the benefits of clonal integration in an invasive plant spreading from open patches into plant communities

• One benefit of clonal integration is that resource translocation between connected ramets enhances the growth of the ramets grown under stressful conditions, but whether such resource translocation reduces the performance of the ramets grown under favourable conditions has not produced consistent results. In this study, we tested the hypothesis that resource translocation to recipient ramets may reduce the performance of donor ramets when resources are limiting but not when resources are abundant.
• We grew Mikania micrantha stolon fragments (each consisting of two ramets, either connected or not connected) under spatially heterogeneous competition conditions such that the developmentally younger, distal ramets were grown in competition with a plant community and the developmentally older, proximal ramets were grown without competition. For half of the stolon fragments, slow‐release fertiliser pellets were applied to both the distal and proximal ramets.
• Under both the low and increased soil nutrient conditions, the biomass, leaf number and stolon length of the distal ramets were higher, and those of the proximal ramets were lower when the stolon internode was intact than when it was severed. For the whole clone, the biomass, leaf number and stolon length did not differ between the two connection treatments. Connection did not change the biomass of the plant communities competing with distal ramets of M. micrantha.
• Although clonal integration may promote the invasion of M. micrantha into plant communities, resource translocation to recipient ramets of M. micrantha will induce a cost to the donor ramets, even when resources are relatively abundant.

     

Metabolite profiles of interacting mycelial fronts differ for pairings of the wood decay basidiomycete fungus, Stereum hirsutum with its competitors Coprinus micaceus and Coprinus disseminatus

The paper presents the first proof-of principle study of metabolite profiles of the interacting mycelial fronts of a wood decomposer basidiomycete, Stereum hirsutum, paired with two competitor basidiomycetes, Coprinus disseminatus and C. micaceus, using TLC and GC-TOF-MS profiling. GC-TOF-MS profiles were information rich, with a total of 190 metabolite peaks detected and more than 120 metabolite peaks detected per sample. The metabolite profiles were able to discriminate between the interactions of S. hirsutum with the two species of Coprinus. In confrontation with C. micaceus, where S. hirsutum mycelial fronts always overgrew those of C. micaceus, there were down-regulations of metabolites in the interaction zone, compared to monocultures of both S. hirsutum and C. micaceus. In contrast, in pairings with C. disseminatus, whose mycelia overgrew those of S. hirsutum, there were some up-regulations compared with monoculture controls, the majority of the metabolites being characteristic of the S. hirsutum monoculture profile. These differences indicate that up-regulation of metabolites in the mycelia of S. hirsutum may be connected to a defensive role or to stress. The results also show proof of principle for the employment of metabolic profiling for biological discovery studies of metabolites produced by fungi that could be applied to natural product screening programmes.     

Non‐trophic effects of oribatid mites on cord‐forming basidiomycetes in soil microcosms

1. Saprotrophic cord‐forming basidiomycetes are the primary agents of decomposition in forest ecosystems. Collembola and oribatid mites affect fungal growth and foraging, and therefore decomposition, through direct mycelial grazing. 2. Grazing on the fungal species Hypholoma fasciculare, Resinicium bicolor and Phanerochaete velutina by the collembola Folsomia candida, and the oribatid mites Steganacarus magnus, Euzetes globulus and Hermannia gibba was investigated in soil microcosms. 3. Folsomia candida grazed on all fungal species: radial extent of R. bicolor, hyphal coverage of all fungal species, and fractal dimension of R. bicolor and P. velutina were all reduced. Oribatid mites did not graze the fungi but did affect mycelial morphology. Steganacarus magnus caused a reduction in the radial extent of H. fasciculare, and the hyphal coverage and fractal dimension in both H. fasciculare and R. bicolor. Euzetes globulus and H. gibba reduced the hyphal coverage of P. velutina. 4. Oribatid mites are associated with a cornucopia of chemical secretions with possible anti‐fungal properties. Chemical analysis of H. gibba opisthonotal secretions revealed four main compounds, all of which are new to the known spectrum of opisthonotal components. The most abundant was (E)‐β‐farnesene. 5. Treatment effects were species‐specific in terms of both fungal and invertebrate species. This study provides the first evidence of non‐grazing effects of oribatid mites on fungal growth and morphology. This could potentially influence the spatial organisation of mycelium in forest soils and therefore the ability of fungi to locate, colonise and decompose dead organic matter.     

Applying dimorphic yeasts as model organisms to study mycelial growth: Part 1. Experimental investigation of the spatio-temporal development of filamentous yeast colonies

Colony development of the dimorphic yeasts Yarrowia lipolytica and Candida boidinii on solid agar substrates under glucose limitation served as a model system for mycelial development of higher filamentous fungi. Strong differences were observed in the behaviour of both yeasts: C. boidinii colonies reached a final colony extension which was small compared to the size of the growth field. They formed cell-density profiles which steeply declined along the colony radius and no biomass decay processes could be detected. The stop of colony extension coincided with the depletion of glucose from the growth substrate. These findings supported the hypothesis that glucose-limited C. boidinii colonies can be regarded as populations of single cells which grow according to a diffusion-limited growth mechanism. Y. lipolytica colonies continued to extend after the depletion of the primary nutrient resource, glucose, until the populations covered the entire growth field which was accomplished by utilization of mycelial biomass.     

Does carbon partitioning in ectomycorrhizal pine seedlings under elevated CO2 vary with fungal species?

Enhanced soil respiration in response to elevated atmospheric CO₂ has been demonstrated, and ectomycorrhizal (ECM) fungi are of particular interest since they partition host-derived photoassimilates belowground. Although a strong response of ECM fungi to elevated CO₂ has been shown, little is still known about the functional diversity among species. We studied carbon (C) partitioning in mycorrhizal Scots pine seedlings in response to short-term CO₂ enrichment, using seven ECM species with different ecological strategies. Mycorrhizal associations were synthesised and seedlings grown in large Petri dishes containing peat:vermiculite and nutrient solution for 10–15 weeks, after which half of the microcosms were exposed to elevated CO₂ treatment (710 ppm) for 15 days and the other half were kept in ambient CO₂ treatment. Partitioning of C was quantified by pulse labelling the seedlings with ¹⁴CO₂ and examining the distribution of labelled assimilates in shoot, root and extraradical mycelial compartments by destructive harvest and liquid scintillation counting. Fungal biomass was determined with PLFA analysis. The respiratory loss of ¹⁴CO₂ was on average greater in the elevated CO₂ treatment for most species compared to the ambient CO₂ treatment. More label was retrieved in the shoots in the ambient CO₂ treatment compared to elevated CO₂ (significant for P. involutus and P. fallax). Greater amounts of label were found in the extraradical mycelial compartment in all species (except P. involutus) in elevated CO₂ compared to ambient CO₂ (significant for L. bicolor, P. byssinum, P. fallax and R. roseolus). Fungal biomass production increased significantly with elevated CO₂ for two species (H. velutipes and A. muscaria); three species (P. fallax, P. involutus and R. roseolus) showed a similar but non-significant trend, whereas L. bicolor and P. byssinum produced less biomass in elevated CO₂ compared to ambient CO₂. When ¹⁴C in the mycelial compartment and respiration was expressed per unit fungal PLFA the difference between CO₂ treatments disappeared. We demonstrated that different ECM fungal isolates respond differently in C partitioning in response to CO₂ enrichment. These results suggest that under certain growth conditions, when nutrients are not limiting, ECM fungi respond rapidly to increasing C-availability through changed biomass production and respiration.     

Manipulation of food resources by a gall-forming aphid: the physiology of sink-source interactions

Summary We examined the capacity of the galling aphid, Pemphigus betae, to manipulate the sink-source translocation patterns of its host, narrowleaf cottonwood (Populus angustifolia). A series of ¹⁴C-labeling experiments and a biomass allocation experiment showed that P. betae galls functioned as physiologic sinks, drawing in resources from surrounding plant sources. Early gall development was dependent on aphid sinks increasing allocation from storage reserves of the stem, and later development of the progeny within the gall was dependent on resources from the galled leaf blade and from neighboring leaves. Regardless of gall position within a leaf, aphids intercepted ¹⁴C exported from the galled leaf (a non-mobilized source). However, only aphid galls at the most basal site of the leaf were strong sinks for ¹⁴C fixed in neighboring leaves (a mobilized source). Drawing resources from neighboring leaves represents active herbivore manipulation of normal host transport patterns. Neighboring leaves supplied 29% of the ¹⁴C accumulating in aphids in basal galls, while only supplying 7% to aphids in distal galls. This additional resource available to aphids in basal galls can account for the 65% increase in progeny produced in basal galls compared to galls located more distally on the leaf and limited to the galled leaf as a food resource. Developing furits also act as skins and compete with aphid-induced sinks for food supply. Aphid success in producing galls was increased 31% when surrounding female catkins were removed.     

Applying dimorphic yeasts as model organisms to study mycelial growth: part 2. Use of mathematical simulations to identify different construction principles in yeast colonies

The dimorphic yeasts Candida boidinii and Yarrowia lipolytica were applied as model organisms to study mycelial growth. A mathematical model of hybrid cellular automaton type was developed to analyze the impact of different biological assumptions on the predicted development of filamentous yeast colonies. The one-dimensional model described discrete cells and continuous distribution of nutrients. The simulation algorithm accounted for proliferation of cells, diffusion of nutrient, as well as biomass decay and recycling inside the mycelium. Simulations reproduced the spatio-temporal development of C. boidinii colonies when a diffusion-limited growth algorithm based on the growth of pseudohyphal cells was applied. Development of Y. lipolytica colonies could only be reproduced when proliferation was restricted to the colony boundary, and cell decay and biomass recycling were incorporated into the model. The results suggested that cytoplasm, which served as the secondary nutrient resource, had to be translocated inside the hyphal network.     

Environmental heterogeneity and clonal growth: a study of the capacity for reciprocal translocation in Glechoma hederacea L.

Clonal fragments of Glechoma hederacea L. (Lamiaceae) were subjected to environments in which light and nutrients were supplied with a strictly negative association in space, i.e. when one of these resources was in ample supply the other was scarce. Treatments were chosen to simulate environments in which clones grew either within homogeneous conditions or across patch types (heterogeneous conditions). The hypothesis was tested that reciprocal translocation (i.e. exchange of both nutrients and assimilates) between connected groups of ramets would increase biomass production of clones growing under heterogeneous conditions compared to that of clones growing in homogeneous conditions. A cost-benefit analysis was carried out to test this hypothesis. Results suggested that reciprocal translocation did not occur at the structural scale considered in this experiment; no evidence was found for a significant effect on whole clone biomass of assimilate and/or nutrient translocation between clone parts experiencing contrasting levels of resource supply. It is suggested that predominantly acropetal movement of resources and the pattern of integrated physiological unit formation in G. hederacea are the main properties responsible for the lack of mutual physiological support between connected clonal fragments growing in differing habitat conditions. These properties are expected to promote clonal expansion and the exploitation of new territory, rather than sustaining clone parts in sub-optimal patches of habitat for prolonged periods of time.     

Verticillium wilt of cacao in Uganda: the relationship between Verticillium dahliae and cacao roots

Isolates of Verticillium dahliae Kleb. from wilted cacao (Theobroma cacao L.), cotton (Gossypium hirsutum L.), and okra (Abelmoschus esculentus Medik.) penetrated all regions of living cacao tap and lateral roots and progressed intracellularly from the epidermis to the xylem in 4–6 days. The hypocotyl and tissues of the unerupted lateral roots beneath the epidermis resisted invasion. Host reactions included browning of extensively colonized cells, alteration (with apparent granulation) of the cytoplasm, and accumulation of materials in the lumina of endodermal cells. Resistance in the hypocotyl was associated with occasional thickening of inner tangential walls of colonized epidermal cells. The fungus formed conidia, microsclerotia, and narrow and wide hyphae within root tissues. The narrow hyphae predominated at the front of mycelial invasion of tissues while the broad hyphae developed behind this front. Limited studies under non-sterile conditions indicated that the apparent host-parasite interactions were similar to those observed with sterile roots and cultures of V. dahliae.     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.     

Resource translocation within seagrass clones: allometric scaling to plant size and productivity

The allometric scaling of resource demand and translocation within seagrass clones to plant size (i.e. shoot mass and rhizome diameter), shoot production and leaf turnover was examined in situ in eight seagrass species (Cymodocea nodosa, Cymodocea serrulata, Halophila stipulacea, Halodule uninervis, Posidonia oceanica, Thalassodendron ciliatum, Thalassia hemprichii and Zostera noltii), encompassing most of the size range present in seagrass flora. One fully developed shoot on each experimental rhizome was incubated for 2–3 h with a pulse of NaH¹³CO₃ (235 μmol) and ¹⁵NH₄Cl (40 μmol). The mobilisation of incorporated tracers across the clone was examined 4 days later. Carbon and nitrogen demand for shoot production across seagrass species scaled at half of the shoot mass, whereas seagrass leaves incorporated tracers (¹³C and ¹⁵N) at rates proportional to the shoot mass. The shoots of all seagrass species shared resources with neighbours, particularly with younger ones. The time scales of physiological integration and the absolute amount of resources shared by seagrass ramets scaled at 2.5 power of the rhizome diameter. Hence, the ramets of larger species were physiologically connected for longer time scales and share larger absolute amounts of resources with neighbours than those of smaller species. The different pattern of resource translocation exhibited by seagrasses helps explain the ecological role displayed by these species and the success of large seagrasses colonising nutrient-poor coastal areas, where they often dominate.     

五种矿质元素对草菇菌丝生长和活性氧清除能力的影响

【背景】菌种退化是草菇生产中面临的难题,探究一种简单高效的复壮方法是草菇产业亟须解决的问题。【目的】探讨外源添加5种矿质元素对草菇退化菌株的菌丝性状和活性氧(reactive oxygen species, ROS)清除能力的影响。【方法】筛选出5种矿质元素的最佳浓度并添加于PDA培养基中,测定草菇菌落形态、菌丝生长速度、菌丝生物量、ROS含量、抗氧化酶活性及抗氧化物质含量。【结果】MnSO4、Na2SeO3、CaSO4、FeSO4可有效提高草菇退化菌株D1和D2的菌丝生长速度和生物量;降低O2·-、H2O2等ROS的含量,并增加还原型谷胱甘肽(glutathione, GSH)、氧化型谷胱甘肽(glutathionedisulfide,GSSG)的含量及超氧化物歧化酶(superoxidedismutase,SOD)、过氧化氢酶(catalase, CAT)、谷胱甘肽...     

Nutrient and water addition effects on day- and night-time conductance and transpiration in a C3 desert annual

Recent research has shown that many C₃ plant species have significant stomatal opening and transpire water at night even in desert habitats. Day-time stomatal regulation is expected to maximize carbon gain and prevent runaway cavitation, but little is known about the effect of soil resource availability on night-time stomatal conductance (g) and transpiration (E). Water (low and high) and nutrients (low and high) were applied factorially during the growing season to naturally occurring seedlings of the annual Helianthus anomalus. Plant height and biomass were greatest in the treatment where both water and nutrients were added, confirming resource limitations in this habitat. Plants from all treatments showed significant night-time g (~0.07 mol m⁻² s⁻¹) and E (~1.5 mol m⁻² s⁻¹). In July, water and nutrient additions had few effects on day- or night-time gas exchange. In August, however, plants in the nutrient addition treatments had lower day-time photosynthesis, g and E, paralleled by lower night-time g and E. Lower predawn water potentials and higher integrated photosynthetic water-use efficiency suggests that the nutrient addition indirectly induced a mild water stress. Thus, soil resources can affect night-time g and E in a manner parallel to day-time, although additional factors may also be involved.     

Population structure and responses to disturbance of the basidiomycete Resinicium bicolor

Summary Resinicium bicolor (Alb. & Schw. ex Fr.) Parm. [=Odontia bicolor (Alb. & Schw. ex Fr.) Bres.] is an outcrossing resupinate basidiomycete associated with root and butt rots of trees, but is itself only very weakly pathogenic. The distribution of genets among every spruce stump in a 70-year-old 1250 m² spruce stand was analysed using somatic incompatibility testing. R. bicolor was present on 40% of 8-to 10-year-old stumps. Nineteen genets were found occupying 32 stumps; yielding probabilities of colonisation following establishment by basidiospores of 0.20–0.24 and by mycelial extension or dispersal of 0.16–0.20. The probability of colonisation decreased with increasing distance from a point of establishment. R. bicolor responded to both enrichment and destructive disturbances by the formation of an extensive cord system which enabled it to colonise discontinuously distributed resources and to overgrow fungi adjacent to it in a single resource unit, including Heterobasidion annosum.     

Impacts of elevated temperature on the growth and functioning of decomposer fungi are influenced by grazing collembola

Elevated temperature has potential to influence the biological mechanisms regulating ecosystem–atmosphere carbon exchange. The relationship between warming and heterotrophic microbial respiration remains poorly understood, not least in terms of the differential sensitivity of microbial groups to temperature and the complexity of interactions with other biota. Cord‐forming basidiomycete fungi are dominant primary decomposers in temperate woodland. Decomposition rates are determined by the composition of the decomposer community, ecophysiological relationships between these fungi and abiotic variables and interactions with other organisms. Amongst the latter, a major determinant is the balance between mycelial growth and removal by soil invertebrate grazers, which can themselves be affected by elevated temperature. We investigated the impact of elevated temperature on fungal foraging and decomposition of beech (Fagus sylvatica) wood in soil microcosms to which the invertebrate grazers, Folsomia candida and Protophorura armata (Collembola), were added in factorial combinations with five basidiomycete fungi. Species‐specific impacts on mycelial development and function resulted from differential sensitivity of fungi to warming and grazing. Temperature impacts on collembola abundance were resource‐specific, causing increased grazing pressure by both species, but on different fungi. Grazing often counteracted warming‐induced stimulation of mycelial growth, but occasionally amplified the temperature effect, with implications for colonization rates of new resources. High grazing pressure did not prevent increased fungal‐mediated decomposition of colonized wood, as fungi utilized more resource‐derived energy to maintain explorative growth. Impacts of elevated temperature on decomposition are likely to depend on local composition of the fungal and invertebrate decomposer community.     

Resource translocation drives 13C fractionation during recovery from disturbance in giant kelp,Macrocystis pyrifera

Resource allocation and translocation are fundamental physiological functions for autotrophs. The mobilization and use of resources drive population dynamics by regulating growth and recovery of individuals, but also influences ecosystem‐level processes such as primary productivity and carbon cycling. This study provides the first observation of translocation‐driven gradients of δ¹³C in macroalgae, a critically important phenomenon recognized in vascular plants for decades. A ~10‰ δ¹³C increase in new giant kelp (Macrocystis pyrifera) fronds relative to mature canopy blades was produced after 5 weeks following a biomass removal experiment, more than twice the variation typical for macroalgae. The observed δ¹³C patterns are consistent with tissue enrichment following resource translocation in vascular plants. The analogous source‐sink relationships and consistent translocation patterns in Macrocystis and vascular plants suggest that translocation of stored resources is critical for structuring productivity and recovery from disturbance in important, habitat‐forming macroalgae such as kelps and fucoids.     

Competition for nitrogen between Pinus sylvestris and ectomycorrhizal fungi generates potential for negative feedback under elevated CO2

We investigated fungal species-specific responses of ectomycorrhizal (ECM) Scots pine (Pinus sylvestris) seedlings on growth and nutrient acquisition together with mycelial development under ambient and elevated CO₂. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor, Suillus bovinus, S. luteus, Piloderma croceum, Paxillus involutus, Boletus badius, or non-mycorrhizal, under ambient, and elevated CO₂ (350 or 700 μl l⁻¹ CO₂); each treatment contained six replicates. The trial lasted 156 days. During the final 28 days, the seedlings were labeled with ¹⁴CO₂. We measured hyphal length, plant biomass, ¹⁴C allocation, and plant nitrogen and phosphorus concentration. Almost all parameters were significantly affected by fungal species and/or CO₂. There were very few significant interactions. Elevated CO₂ decreased shoot-to-root ratio, most strongly so in species with the largest extraradical mycelium. Under elevated CO₂, ECM root growth increased significantly more than hyphal growth. Extraradical hyphal length was significantly negatively correlated with shoot biomass, shoot N content, and total plant N uptake. Root dry weight was significantly negatively correlated with root N and P concentration. Fungal sink strength for N strongly affected plant growth through N immobilization. Mycorrhizal fungal-induced progressive nitrogen limitation (PNL) has the potential to generate negative feedback with plant growth under elevated CO₂. Responsible Editor: Herbert Johannes Kronzucker     

UPTAKE AND TRANSLOCATION OF TI FROM NANOPARTICLES IN CROPS AND WETLAND PLANTS

Bioavailability of engineered metal nanoparticles affects uptake in plants, impacts on ecosystems, and phytoremediation. We studied uptake and translocation of Ti in plants when the main source of this metal was TiO₂ nanoparticles. Two crops (Phaseolus vulgaris (bean) and Triticum aestivum (wheat)), a wetland species (Rumex crispus, curly dock), and the floating aquatic plant (Elodea canadensis, Canadian waterweed), were grown in nutrient solutions with TiO₂ nanoparticles (0, 6, 18 mmol Ti L^?1 for P. vulgaris, T. aestivum, and R. crispus; and 0 and 12 mmol Ti L^?1 for E. canadensis). Also examined in E. canadensis was the influence of TiO₂ nanoparticles upon the uptake of Fe, Mn, and Mg, and the influence of P on Ti uptake. For the rooted plants, exposure to TiO₂ nanoparticles did not affect biomass production, but significantly increased root Ti sorption and uptake. R. crispus showed translocation of Ti into the shoots. E. canadensis also showed significant uptake of Ti, P in the nutrient solution significantly decreased Ti uptake, and the uptake patterns of Mn and Mg were altered. Ti from nano-Ti was bioavailable to plants, thus showing the potential for cycling in ecosystems and for phytoremediation, particularly where water is the main carrier.     

Translocation,remineralization, and turnover of nitrogen in the roots and rhizomes of Spartina alterniflora (Gramineae)

We used ¹⁵N to quantify rates of N translocation from aerial to belowground tissues, foliar leaching, and turnover and production of root and rhizome biomass in the plant-sediment system of short Spartina alterniflora areas of Great Sippewissett Marsh, Massachusetts. Decay of belowground tissues in litterbag incubations at 1- and 10-cm depths resulted in 80% remineralization of the original plant (¹⁵N-labeled) N and 20% burial after 3 years. Translocation of ¹⁵N from plant shoots in hydrologically controlled laboratory lysimeters maintained under field conditions was 38% of the aboveground pool while leaching of N was 10% from June to October. Most of the translocated N was not retranslocated to new aboveground growth in December but appeared to be either remineralized or buried in the sediment. Injection of ¹⁵N into field stands of grass showed initially high incorporation into plants followed by a continuous decline over the next 7 years yielding a gross tumover time of 1.5–1.6yr. Correcting the gross N turnover for recycling of label via translocation and uptake of remineralized label during this period, a net root and rhizome turnover time of 1.0–1.1 yr was obtained. Combining the turnover time with independent estimates of seasonal belowground biomass yielded an estimate of belowground production of 929–1,022 g C m⁻² yr⁻¹, similar to measurements by traditional biomass harvest, CO² based budgets and models for comparable areas of this marsh. Integration of the production and nitrogen balance estimates for short Spartina marsh yielded translocation, 1.4 g N m⁻² yr⁻¹, leaching, 0.4 g N m⁻² yr⁻¹, remineralization, 14.9–16.3 g N m⁻² yr⁻¹, and burial, 3.7–4.1 g N m⁻² yr⁻¹.