Influence of summer sowing dates, N fertilization and irrigation on autumn VSP accumulation and dynamics of spring regrowth in alfalfa (Medicago sativa L.).

Herbage yield of alfalfa (Medicago sativa L.) depends on forage management or environmental conditions that change C and N resource acquisition, and endogenous plants factors such as root organic reserves and number of active meristems. The aim of this work is to study the influence of two sowing dates in summer (12 July or 9 August), N fertilization (0 or 100 kg ha(-1)) and/or irrigation applied during the first year of alfalfa establishment on (i) the accumulation of N organic reserves (soluble proteins and more specifically vegetative storage protein) in taproots during autumn, (ii) the number of crown axillary meristems present at the end of winter and (iii) the dynamics of spring shoot growth. Delaying the sowing date for one month reduced root growth and root N storage, especially vegetative storage proteins (VSP) during autumn. Irrespective of sowing dates, N fertilization did not affect root biomass, number of crown buds, total root N, root soluble protein or VSP concentrations. By contrast, water deficiency during alfalfa establishment in the early summer reduced both root growth and N reserve accumulation. When spring growth resumed, there is a significant linear relationship between leaf area development and soluble protein and VSP concentrations in taproots, and also the number of crown buds. The results showed that an early sowing date and adequate water status during the summer allowed alfalfa plants to accumulate N reserves by increasing taproot mass and soluble protein concentrations, especially VSPs. This resulted in rapid shoot regrowth rates the following spring.     

A role for nitrogen reserves in forage regrowth and stress tolerance

Carbohydrate accumulation and utilization during shoot regrowth after defoliation and winter has been studied extensively in most species used as forage. However, recent work suggests that N reserves found in vegetative tissues also are important for defoliation tolerance and winter hardiness. Results suggest that these N reserves constitute an alternative N source used when N₂ fixation and/or mineral N uptake are reduced. ¹⁵N labelling experiments indicate that a large proportion of herbage N is derived from N reserves mobilized from stem bases or roots to developing leaves and shoots. Amino acids and specific proteins (i.e. vegetative storage proteins, VSPs) are deposited in roots and stem bases and, in the case of VSPs, are degraded rapidly after defoliation. Identification and characterization of VSPs will increase our understanding of the role N reserves play in stress tolerance and may lead to innovative approaches for improving forage persistence and productivity.     

Mobilization of nitrogen reserves during regrowth of defoliated Trifolium repens L. and identification of potential vegetative storage proteins

Although it is well established that carbon reserves contributeto shoot regrowth of leguminous forage species, little informationis available on nitrogen reserves except in Medicaqo sativaL. and Trifolium subterraneum L. In this study, reserves werelabelled with ¹⁵N to demonstrate the mobilization of endogenousnitrogen from roots and stolons to regrowing leaves and newstolons during 24 d of regrowth in white clover (Thfolium repensL.). About 55% and 70%, respectively, of the nitrogen contentsof these organs were mobilized to support the regrowth of leaves.During the first 6 d, nitrogen in regrowing leaves came mainlyfrom N reserves of organs remaining after defoliation. Afterthese first 6 d of regrowth, most of the shoot nitrogen wasderived from exogenous nitrogen taken up while the contributionof nitrogen reserves decreased. After defoliation, the buffer-solubleprotein content of roots and stolons decreased by 32% duringthe first 6 d of regrowth. To identify putative vegetative storageproteins, soluble proteins were separated using SDS-PAGE ortwo-dimensional electrophoresis. One protein of 17.3 kDa instolons and two proteins of 15 kDa in roots seemed to behaveas vegetative storage proteins. These three polypeptides, initiallyfound at high concentrations, decreased in relative abundanceto a large extent during early regrowth and then were accumulatedagain in roots and stolons once normal growth was re-established. Key words: White clover, regrowth, ¹⁵N-labelled, vegetative storage proteins, electrophoresis     

Potato leafhopper (Homoptera: Cicadellidae) injury disrupts basal transport of 14C-labelled photoassimilates in alfalfa

The potato leafhopper, Empoasca fabae (Harris), is a key pest of alfalfa, Medicago sativa L., in part because of the leafhopper's ability to disrupt upward translocation within phloem tissues. To determine if leafhopper injury also disrupts basal translocation necessary for regrowth and perenniality of alfalfa, we used radiolabeled 14CO2 to measure the basal transport of photoassimilates in injured and healthy plants. In one experiment, less 14C was transported to lower stem tissue of leafhopper-injured plants in comparison to the same tissue of healthy plants in early vegetative and early reproductive stages of alfalfa development. In a second experiment, less 14C was transported to lower stem, crown, and root tissues of injured plants in comparison to the same tissues of healthy, early reproductive plants. The disruption of basal transport caused by potato leafhopper may impact carbon storage and mobilization subsequent to defoliation, winter survival, and nitrogen fixation.     

The use of internal nitrogen stores in the rhizomatous grass Calamagrostis epigejos during regrowth after defoliation

BACKGROUND AND AIMS: The regrowth dynamics after defoliation of the invasive grass Calamagrostis epigejos were studied. As nitrogen (N) reserves have been shown to play an important role during plant regrowth, the identity, location and relative importance for regrowth of N stores were determined in this rhizomatous grass. METHODS: Plant growth, nitrate uptake and root respiration were followed during recovery from defoliation. Water soluble carbohydrates, nitrate, free amino acids and soluble proteins were analysed in the remaining organs. KEY RESULTS: Nitrate uptake and root respiration were severely reduced during the first days of regrowth. Roots were the main net source of mobilized N. The quantitatively dominant N storage compounds were free amino acids. Free amino acids and soluble proteins in the roots decreased by 55 and 50%, respectively, and a substantial (approximately 38%) decrease in stubble protein was also observed. Although the relative abundance of several soluble proteins in roots decreased during the initial recovery from defoliation, no evidence was found for vegetative storage protein (VSP). Furthermore, rhizomes did not act as a N storage compartment. CONCLUSIONS: Production of new leaf area was entirely reliant, during the first week after defoliation, on N stores present in the plant. Mobilized N originated mainly from free amino acids and soluble proteins located in roots, and less so from proteins in stubble. Presence of VSP in the roots was not confirmed. The data suggest that rhizomes played an important role in N transport but not in N storage.     

Nutrient reserves in roots of fruit trees,in particular carbohydrates and nitrogen

Summary In trees, nutrient reserves built up in the previous year are of primary importance for early spring growth. Despite the relatively great importance of roots for nutrient storage, the root system should not be regarded as a special storage organ. Quantitatively, carbohydrates predominate in these reserves, but qualitatively N and other minerals are of more than minor significance. In roots carbohydrates are usually stored in insoluble form, mainly as starch; sorbitol is the predominant soluble compound in apple and peach. For nitrogen reserves, the soluble form predominates in roots, especially arginine in apple and peach, followed by asparagine. The level of reserves usually becomes maximal early in the winter. During leafing-out the reserves are drawn on until, later in the season, the supply of newly produced or absorbed nutrients exceeds the demand and replenishment occurs. The initial carbohydrate reserves do not determine the amount of new growth, whereas reserve nitrogen is of decisive importance for shoot growth vigour. Environmental factors such as light intensity and temperature affect the level of carbohydrates in roots; the concentration can be reduced by defoliation and summer pruning and increased by ample supply of nitrogen fertilizer in the autumn. The main cultural factors that influence nitrogen reserves are the amount and the time of nitrogen fertilization.     

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.     

Carbohydrate Metabolism in Taproots of Medicago sativa L. during Winter Adaptation and Spring Regrowth

Our objective was to identify amylases that may participate in starch degradation in alfalfa (Medicago sativa L.) taproots during winter hardening and subsequent spring regrowth. Taproots from field-grown plants were sampled at intervals throughout fall, winter, and early spring. In experiment 1, taproots were separated into bark and wood tissues. Concentrations of soluble sugars, starch, and buffer-soluble proteins and activities of endo- and exoamylase were determined. Starch concentrations declined in late fall, whereas concentrations of sucrose increased. Total amylolytic activity (primarily exoamylase) was not consistently associated with starch degradation but followed trends in soluble protein concentration of taproots. This was especially evident in spring when both declined as starch degradation increased and shoot growth resumed. Activity of endoamylase increased during periods of starch degradation, especially in bark tissues. In experiment 2, a low starch line had higher specific activity of taproot amylases. This line depleted its taproot starch by late winter, after which taproot sugar concentrations declined. As in experiment 1, total amylolytic activity declined in spring in both lines, whereas that of endoamylase increased in both lines even though little starch remained in taproots of the low starch line. Several isoforms of both amylases were distinguished using native polyacrylamide electrophoresis, with isoforms being similar in bark and wood tissues. The slowest migrating isoform of endoamylase was most prominent at each sampling. Activity of all endoamylase isoforms increased during winter adaptation and in spring when shoot growth resumed. Endoamylase activity consistently increased at times of starch utilization in alfalfa taproots (hardening, spring regrowth, after defoliation), indicating that it may serve an important role in starch degradation.     

Carbohydrate storage and use in an alpine population of the perennial herb, Oxytropis sericea

I tested hypotheses for ecological roles of storage carbohydrates in perennating organs (roots and branches) of alpine Oxytropis sericea, a leguminous herb. In naturally growing plants, total nonstructural carbohydrates achieved their maximal concentration in the fall, declined during winter, and reached minimal levels immediately after growth initiation in the spring. Experimental manipulation of carbon sink-source relations through shading of leaves of reproductive plants revealed that the normally unused portion of these carbohydrates is largely available for withdrawal. In another experiment, plants subjected to carbohydrate depletion through shading suffered decreased leaf growth after winter dormancy and had a lower probability of flowering and decreased inflorescence biomass. The dependence of reproductive growth on stored carbohydrates, however, was limited to its initial stages, because accumulation of storage carbohydrates occurred simultaneously with inflorescence expansion, flowering, and fruiting. Moreover, the whole-plant photosynthetic rate, estimated from gas exchange measurements also peaked at the time of inflorescence growth. To address whether stored reserves allow compensatory regrowth following defoliation, plants were subjected to experimental removal of leaves and inflorescences. Defoliated O. sericea partly regrew the lost leaves but withdrawal of stored carbohydrates was limited. Similarly, in a second defoliation experiment where infructescences were left intact, the plants used little stored carbohydrate and only partly compensated for fruit growth. However, carbohydrate accumulation was negatively affected by defoliation. While the ecological importance of stored nonstructural carbohydrates cannot be attributed to any function in isolation, winter respiration, leaf regrowth after winter, and early reproductive growth in O. sericea all depend to a significant extent on stored reserves. Maintaining a large storage pool may protect these functions in years when carbon status is less favorable than during this study. Received: 13 May 1998 / Accepted: 24 November 1998     

Effects of aboveground herbivory on plants with long-term belowground biomass storage

Plant tolerance to herbivory is contingent on multiple traits and adaptive mechanisms, which makes it a complex response with ecological implications. In plants with long-term belowground storage, allocation of biomass to inaccessible parts belowground in response to folivory is a well-recognized tolerance mechanism. In temperate regions, spring growth from buried rootstock is common among winter deciduous plants and is often followed by regrowth after defoliation, both of which draws resources from the stored reserves. We developed a mathematical model to analyze this tolerance response in a winter deciduous plant with long-term belowground biomass when it is defoliated by a specialist insect folivore. The model explores how three closely associated traits—(1) belowground biomass allocation to roots, (2) spring utilization of stored reserves, and (3) post-defoliation regrowth capacity—modulate the persistence and dynamics of the plant and herbivore populations. Model results show that allocation to belowground storage is not only a critical component of tolerance but also influences the herbivore population dynamics in ways that depend on how and when plant biomass is allocated and used. Low belowground biomass allocation and high storage utilization combined with poor photosynthetic growth caused extirpation of the plant population by the defoliating insects. Stable coexistence of the plant at low biomass along with its specialist insect required a moderate amount of post-herbivory belowground allocation. High values of belowground biomass allocation, storage utilization, and photosynthetic growth resulted in sustained cycles of the herbivore and plant populations. Interestingly, utilization of stored reserves had conflicting influence on above and belowground biomass, and strongly affected herbivore population dynamics. Our model thus highlights the complexity of tolerance response when it involves multiple traits and mechanisms as evinced by winter deciduous plants. We close by discussing the implications of our findings for the contributions of defoliating insects to biocontrol programs.     

沙棘根瘤内生菌的多型性

用透射电子显微镜观察了春、夏、秋、冬四个季节的沙棘根瘤,以及瘤瓣上、中、下三个部位。结果表明,不同季节,不同部位的瘤瓣内,根瘤内生菌有7种不同形态。即侵染菌丝体、繁殖菌丝体、营养菌丝体、春孢子及春孢子囊、泡囊,冬孢子及冬孢子囊和类菌体。在多年生珊瑚状的根瘤中,它们的世代交替是:春夏季以侵染菌丝、繁殖菌丝、营养菌丝、春孢子囊及春孢子、泡囊为主;秋冬季以衰退的营养菌丝、衰老泡囊、冬孢子囊和冬孢子、类菌体为主。冬孢子和类菌体是休眠体。     

Morphological pattern of development affects the contribution of nitrogen reserves to regrowth of defoliated white clover (Trifolium repens L.)

The contribution of nitrogen reserves to regrowth following defoliation was studied in white clover plants (Trifolium repens cv. Huia). This was found to be closely linked to the morphological pattern of development of the aerial parts during the same period. Low temperature (6 degrees C) and short day exposure (8 h photoperiod) were used to induce dwarf development, i.e. to increase branching rate and to enhance new sites of leaf production during a period of regrowth. Treated plants exhibited a large reduction in leaf area and a large increase in leaf pool size for the first 10 d of a subsequent regrowth under standard culture conditions (16 h daylight; 22/18 degrees C day/night). The contribution of nitrogen from storage compounds in organs remaining after defoliation (sources) to regrowing tissues (sinks) was assessed by 15N pulse-chase labelling during regrowth following shoot removal. The mobilization of nitrogen reserves from storage tissues of regrowing clover was closely linked to the pattern of differentiation of the newly developed organs. It appeared that regrowth was supported less by endogenous N for the first 10 d after defoliation in treated plants, compared with control plants grown continuously in standard conditions. It is assumed that dwarf plants exhibit a lower dependence upon the mobilization of soluble proteins previously accumulated in roots and uncut stolons. The relationship between leaf development rate and N-uptake recovery following defoliation is discussed.     

Effect of elevated CO2, temperature and drought on dry matter partitioning and photosynthesis before and after cutting of nodulated alfalfa

The rising atmospheric CO₂ concentration resulting from industrial development may enhance photosynthesis and plant growth. However, there is a lack of research concerning the effect of combined factors such as CO₂, temperature and water availability on plant regrowth following cutting or grazing, which represent the usual methods of managing forage legumes like alfalfa. Elevated CO₂, temperature and drought can interact with cutting factors (e.g. cutting frequency or height), and source-sink balance differences before and after defoliation can modify photosynthetic behaviour and dry matter accumulation, as well as dry matter partitioning between above- and belowground organs. The aim of our study was to determine the interactive effect of CO₂ (ambient, around 350 μmol mol⁻¹ versus 700 μmol mol⁻¹), temperature (ambient versus ambient + 4 °C) and water availability (well-irrigated versus partially irrigated) on dry matter partitioning and photosynthesis in nodulated alfalfa after vegetative normal growth and during regrowth. At the end of vegetative normal growth, CO₂ enhanced dry matter accumulation despite photosynthesis being down-regulated at the end of this period. Photosynthesis was stimulated by elevated CO₂ and resulted in greater dry matter accumulation during the regrowth period. Aboveground organs were affected more by drought than belowground organs during the entire experiment, particularly during vegetative normal growth. The higher drought tolerance (greater growth) observed during the regrowth period may be related to higher mass and greater reserves accumulated in the roots of plants.     

Interspecific and intraspecific variation in tree seedling survival: effects of allocation to roots versus carbohydrate reserves

We examined interspecific and intraspecific variation in tree seedling survival as a function of allocation to carbohydrate reserves and structural root biomass. We predicted that allocation to carbohydrate reserves would vary as a function of the phenology of shoot growth, because of a hypothesized tradeoff between aboveground growth and carbohydrate storage. Intraspecific variation in levels of carbohydrate reserves was induced through experimental defoliation of naturally occurring, 2-year-old seedlings of four northeastern tree species –Acer rubrum, A. saccharum, Quercus rubra, and Prunus serotina– with shoot growth strategies that ranged from highly determinate to indeterminate. Allocation to root structural biomass varied among species and as a function of light, but did not respond to the defoliation treatments. Allocation to carbohydrate reserves varied among species, and the two species with the most determinate shoot growth patterns had the highest total mass of carbohydrate reserves, but not the highest concentrations. Both the total mass and concentrations of carbohydrate reserves were significantly reduced by defoliation. Seedling survival during the year following the defoliation treatments did not vary among species, but did vary dramatically in response to defoliation. In general, there was an approximately linear relationship between carbohydrate reserves and subsequent survival, but no clear relationship between allocation to root structural biomass and subsequent survival. Because of the disproportionate amounts of reserves stored in roots, we would have erroneously concluded that allocation to roots was significantly and positively related to seedling survival if we had failed to distinguish between reserves and structural biomass in roots. Received: 14 December 1999 / Accepted: 2 June 1999     

SEASONAL PATTERNS IN RECOVERABLE UNDERGROUND RESERVES IN SPARTINA ALTERNIFLORA LOISEL.

Seasonal differences in recoverable underground reserves in stands of Spartina alterniflora Loisel. were measured in Georgia and Delaware. They indicate that by winter substantial reserves are stored in both northern and southern marshes. Cycles of the quantity of stored reserves have greater amplitudes than cycles of the concentration of storage compounds reported in the literature. Maximum concentrations apparently occur at the same time as maximum storage tissue volumes. Seasonal measurements and defoliation studies in Delaware indicate that reserves are nearly exhausted by spring growth and that at least ⅓ are not replenished until after seed filling in the fall. This pattern of nearly complete utilization of reserves in spring with late fall replenishment may have an advantage for herbaceous perennial plants which are part of detritus based food webs. A cycle where substantial reserves are retained for canopy regeneration after early summer defoliation would be advantageous where a grazing web was dominant.     

Nitrogen Reserve Mobilization during Regrowth of Medicago sativa L. (Relationships between Availability and Regrowth Yield)

An experiment was designed to study the role of N and C reserves on regrowth of the shoots following defoliation of forage species. Starch and N accumulation in root and crown tissue of nonnodulated Medicago sativa L. were modified during regrowth by applying different levels of N and different cutting heights. Plants were obtained with similar crown and root dry weights, but having either low starch and high tissue N or high starch and low tissue N. The plants were then submitted to a second defoliation and supplied with optimal N nutrition, and N flow from reserve was quantified using pulse-chase 15N labeling. Maximum yields following the second regrowth were obtained from those plants having a high tissue N, despite their low level of nonstructural carbohydrate. When N in the roots and crown exceeded 5 mg N plant-1 at the beginning of regrowth, about 68% was translocated to regrowing shoots. Highly significant correlations were also found between the amounts of N available in roots and crown at the beginning of regrowth and (a) the amount of N that was mobilized to new tissues, (b) the amount of N taken up during the regrowth period, and (c) the final shoot yield after 24 d of regrowth. No similar correlations were found for plants that varied in their initial starch content of roots and crown. It is suggested that N reserves were used mainly during the first 10 d after defoliation, and that the resulting aerial growth during this period should be sufficient to restore N2 fixation and/or N uptake to levels equal to those prior to defoliation. These data emphasize (a) the importance of root N reserves in initiating and sustaining new shoot growth, and (b) the need for a re-evaluation of the contribution of C reserves to shoot regrowth.     

The consequences of lower nitrogen availability in autumn for internal nitrogen reserves and spring growth of <Emphasis Type="Italic">Calamagrostis epigejos</Emphasis>

The building and use of internal N stores in the grass Calamagrostis epigejos was investigated in context of complex ecological study focused on mechanisms underlying competitive ability of this highly successful invasive species. Induced changes in nitrogen availability in the course of two subsequent vegetation seasons were used as a tool for finding (i) to what extent high N availability in substrate is important for building N reserves in autumn that support spring regrowth and, (ii) if contrasting contents of N storage compounds may result in differences in growth in the next season. Plants were grown in solely inorganic substrate and received a nutrient solution containing 5 mol m⁻³ of NH₄NO₃. The nitrogen supply was reduced in a low nitrogen (LN) treatment to 0.25 mol m⁻³ in August whereas in high nitrogen (HN) treatment remained high till December. During the following growing season were plants from both treatments grown at the low N supply (0.25 mol m⁻³). An increase in the content of N storage compounds was observed from September to December in both treatments. Plants in the LN treatment showed significantly lower total N content and also N allocated to mobilizable reserves (20–50% of HN plants), namely due to a smaller accumulation of amino acids and soluble protein in autumn. External nitrogen availability in autumn is hence highly important for building N reserves in this species. A major portion of the nitrogen stored in HN plants during winter was taken up from growth medium in late autumn, whereas translocation from senescing shoots dominated in LN treatment. During the winter about 50% of N in plants was permanently present in shoots bearing several frost resistant green leaves. Spring regrowth was accompanied by a fast decrease of both total N and the content of N storage compounds in both treatments. Amino acids were identified as the most prominent source of mobilizable N during spring regrowth. Development of leaf area in LN plants was significantly slower in March and April than in HN plants namely due to smaller number of tillers and green leaves per plant. Low N availability in autumn, therefore, may result in restrictions of plant growth and development in the following season.     

Utilization, diet and diet selection by brent geese Branta bernicla bernicla on salt-marshes in Norfolk

The diet and utilization by brent geese of two plant communities ( Limonium/Armeria marsh—a short sward community in the upper mid salt-marsh and composed of several species, and Salicornia marsh—a community of the lower salt-marsh dominated by Salicornia europuea agg. and Aster tripolium ) were studied on the north Norfolk coast. The biomass on these communities was highest in autumn and declined markedly in November owing to the senescence and die-off of the succulent species. This decline was not significantly precipitated by the grazing of the geese but coincided with their switch to inland habitats where the biomass m^-2 and total food available on fields of winter wheat and grassland within the population's home range was 20–30 times greater. Brent geese continued to graze salt-marsh at a low intensity through the winter, feeding largely on Puccinellia maritima. In spring they returned largely to Limonium/Armeria marsh where they significantly reduced the regrowth of Puccinellia maritima.
The main plant species ealen was Puccinellia maritima , especially in mid winter. Salicornia maritima agg. leaves and seeds formed much of the diet in autumn, whereas Triglochin maritima, Plantago maritima and Aster tripolium were important foods on Limonium/Armeria marsh in spring. All these species were selected at some part of the year. Selection was related to the proline (an imino acid used by salt-marsh plants as an osmoregulator) and chloride content and avoidance related to tannins. Two species which contained condensed tannins. Limonium vulgare and Armeria maritima , were rarely eaten. As well as being a nutrient, proline may help to neutralize the effects of tannins in the diet if incorporated into proline-rich proteins.     

A non-targeted metabolomics approach to quantifying differences in root storage between fast- and slow-growing plants

Life history theory posits that slower-growing species should invest proportionally more resources to storage, structural (e.g. stems) or defence traits than fast-growing species. Previously, we showed that the slower-growing monocarpic plants had lower mortality rates and higher bolting probabilities after two defoliation events. Here, we consider a mechanistic explanation, that the slower-growing species invested relatively more resources to storage. We compared the relative levels of root storage compounds between eight monocarpic species using metabolomic profiling, and characterized plant growth using a size-corrected estimate of relative growth rate (RGR). Growth rate was negatively correlated with the proportional allocation of root metabolites identified as sucrose, raffinose and stachyose and with amino acids known for their roles in nitrogen storage, particularly proline and arginine. The total amount and concentration of energy-corrected carbohydrates were also negatively correlated with RGR. Our results show for the first time that slower-growing species invest proportionally more of their total root metabolites in carbon- and nitrogen-storage compounds. We conclude that the increased investment in these reserves is an important resource allocation strategy underlying the growth-survival trade-off in plants.