Dynamics of Fructan and Sucrose Biosynthesis in Crested Wheatgrass

Fructan metabolism occurs in many temperate grasses when ambienttemperatures are cool. Fructan is believed to influence thecold hardiness of grasses and to have osmoregulatory and carbohydratestorage functions. In the cold deserts of the western UnitedStates, crested wheatgrass (Agropyron spp.) usually maintainsmetabolically active leaves throughout the winter. The investigationreported here concerns nonstructural carbohydrate content ofcrested wheatgrass during a growing season (September-June).Fructan accumulated in plant parts during the fall and springwhen temperatures were near 15?C. However, fructan content waslowest when temperatures were coldest. Surose content peakedwhen fructan levels were low and when maximum daily temperaturesremained below 0?C. We propose that fructan biosynthesis inleaf vacuoles provides a sink for photosynthates when temperaturesare too cold for translocation, and growth. Stored fructan thenprovide a readily available substrate for sucrose synthesisand plant growth. Because crested wheatgrass maintains greenleaves during the winter and is capable of photosynthesis attemperatures near 0?C, less stored carbohydrates are requiredfor respiratory maintenance and spring growth than for warm-seasongrasses. ¹Cooperative investigations of the USDA-Agricultural ResearchService and the Utah Agricultural Experiment Station, Logan,Utah 84322, U.S.A. Approved as Journal Paper No. 3755. ²Plant Physiologists, USDA-ARS-Forage and Range Research, Logan,UT 84322-6300, U.S.A. (Received March 25, 1988; Accepted July 11, 1988)     

Differential Effects of Pratylenchus neglectus Populations on Single and Interplantings of Alfalfa Grasses

The invasion by three different Utah populations of Pratylenchus neglectus (UTI, UT2, UT3) was similar in single and interplantings of ''Lahontan'' alfalfa and ''Fairway'' crested wheatgrass at 24 ñ 3 °C. Population UT3 was more pathogenic than UT1 and UT2 on both alfalfa and crested wheatgrass. Inoculum density was positively correlated with an invasion by P. neglectus. Invasions by UT3 at all initial populations (Pi) exceeded that of UT1 and UT2 for both single and interplanted treatments. The greatest reductions in shoot and root weights of alfalfa and crested wheatgrass were at a Pi of 8 P. neglectus/cm³ soil. Pi was negatively correlated with alfalfa and crested wheatgrass shoot and root growth and nematode reproduction. The reproductive factor (Rf) for UT3 exceeded that of UT1 and UT2 in single and interplantings at all inoculum levels. There were no differences in Rfin the Utah populations in single or interplantings. A nematode invasion increased with temperature and was greatest at 30 °C. Population UT3 was more pathogenic than UT1 and UT2 and reduced shoot and root growth at all soil temperatures. Populations UT1 and UT2 reduced shoot and root growth at 20-30 °C. Soil temperature was negatively correlated with shoot and root growth and positively correlated with nematode reproduction. Reproduction of UT3 exceeded that of UT1 and UT2 at all soil temperatures.     

Regrowth of western wheatgrass utilizing 14C-labeled assimilates stored in belowground parts

Summary Results of this study showed that carbohydrates stored in the roots of western wheatgrass are utilized for regrowth following clipping of the aboveground foliage. Shoots remained dependent on carbohydrates stored in roots until sufficient photosynthetic leaf surface was developed to supply carbon to the shoots. During early phenophases, the partitioning of carbohydrates between shoots and roots was identical, indicating equal metabolic demands for carbon from both the shoot and root systems. Subsequent fluctuations in root and shoot carbohydrates may be caused by selected pressure imposed on either the root or shoot systems by physiological changes in these organs.Respiratory losses of ¹⁴C were slower during the early phenophases which may indicate that either the rate of respiration was slower or that recently assimilated nonlabeled carbon sources were utilized for respiration instead of the endogenous sources assimilated earlier in the growth period. re]19760427     

Morphological Analysis of Tillering in Agropyron spicatum and Agropyron desertorum

The caespitose grasses Agropyron spicatum and Agropyron desertorumexhibit a striking difference in tillering response followingexperimental clipping treatment, with plants of A. desertorumproducing up to 18 times more tillers. The two species are similarin many aspects of their phenology and physiology. Previousexamination of current photosynthate production and levels ofstored carbohydrates indicate only slight differences betweenthe species. The possible role of three anatomical/morphologicalconstraints in controlling tillering was examined. No evidencefor such constraints was found. A basal cluster of buds is presenton the parent tillers. The mean bud number per tiller was similarfor both species and the range (3–9) was identical. Nearlyall of the bud apical meristems appeared anatomically viablethroughout the growing season and vascular development occurredto within 250 to 490 µm of the various bud apices of bothspecies. Both normal fall tillers and summer tillers producedunder clipping treatment originated from the largest, most distalbuds of the basal cluster of buds. However, precocious, morphologicallydistinctive, second-order tillers occasionally grew out fromthe smaller, most basal buds of some elongating fall tillers. Agropyron spicatum, Agropyron desertorum, bluebunch wheatgrass, crested wheatgrass, bud, tiller, tillering ability, meristematic potential, vascular development, regrowth     

The invasive annual cheatgrass releases more nitrogen than crested wheatgrass through root exudation and senescence

Plant-soil feedbacks are an important aspect of invasive species success. One type of feedback is alteration of soil nutrient cycling. Cheatgrass invasion in the western USA is associated with increases in plant-available nitrogen (N), but the mechanism for this has not been elucidated. We labeled cheatgrass and crested wheatgrass, a common perennial grass in western rangelands, with ¹⁵N-urea to determine if differences in root exudates and turnover could be a mechanism for increases in soil N. Mesocosms containing plants were either kept moist, or dried out during the final 10 days to determine the role of senescence in root N release. Soil N transformation rates were determined using ¹⁵N pool dilution. After 75 days of growth, cheatgrass accumulated 30 % more total soil N and organic carbon than crested wheatgrass. Cheatgrass roots released twice as much N as crested wheatgrass roots (0.11 vs. 0.05 mg N kg^?1 soil day^?1) in both soil moisture treatments. This occurred despite lower root abundance (7.0 vs. 17.3 g dry root kg^?1 soil) and N concentration (6.0 vs. 7.6 g N kg^?1 root) in cheatgrass vs. crested wheatgrass. We propose that increases in soil N pool sizes and transformation rates under cheatgrass are caused by higher rates of root exudation or release of organic matter containing relatively large amounts of labile N. Our results provide the first evidence for the underlying mechanism by which the invasive annual cheatgrass increases N availability and establishes positive plant-soil feedbacks that promote its success in western rangelands.     

Soil and plant water relations in a crested wheatgrass pasture: response to spring grazing by cattle

Summary Few field studies have attempted to relate effects of actual livestock grazing on soil and plant water status. The present study was initiated to determine the effects of periodic defoliations by cattle during spring on soil moisture and plant water status in a crested wheatgrass (Agropyron cristatum (L.) Gaertn. and A. desertorum (Fisch. ex Link) Schult.) pasture in central Utah. Soil moisture in the top 130 cm of the soil profile was depleted more rapidly in ungrazed plots than in grazed plots during spring and early summer. Soil moisture depletion was more rapid in grazed plots in one paddock after 1 July due to differential regrowth, but there was no difference in soil water depletion between plots in another paddock during the same period. This difference in soil water depletion between paddocks was related to a difference in date of grazing. Although more water had been extracted from the 60 cm to 130 cm depths in ungrazed plots by late September, cumulative soil moisture depletion over the entire 193 cm profile was similar in grazed and ungrazed plots. Prior to 1 July, grazing had no effect on predawn leaf water potentials as estimated by a pressure chamber technique; however, after 1 July, predawn leaf water potentials were lower for ungrazed plants. Midday leaf water potentials were lower for grazed plants before 1 July, but did not differ between grazed and ungrazed plants after 1 July. A 4- to 8-day difference in date of defoliation did not affect either predawn or midday leaf water potentials. The observed differences in water use patterns during spring and early-summer may be important in influencing growth and competitive interactions in crested wheatgrass communities that are subject to grazing by domestic livestock.     

Factors Affecting Population Trends of Plant-Parasitic Nematodes on Rangeland Grasses

The effects of environmental conditions on population trends of plant-parasitic nematodes were studied in experimental plots of five wheatgrasses in the western Utah desert. In a 3-year (1984-86) field study, soil water and temperature affected the population trends of the ectoparasites, Tylenchorhynchus acutoides and Xiphinema americanum, and the migratory endoparasite, Pratylenchus neglectus, on Fairway crested wheatgrass, Agropyron cristatum; ''Hycrest'' crested wheatgrass, A. cristatum X A. desertorura; ''Rosana'' western wheatgrass, Pascopyrum smithii; ''Oahe'' intermediate wheatgrass, Thinopyrum intermedium; and RS-1 hybrid (Elytrigia repens X Pseudoroegneria spicata). The largest soil populations of these nematode species were collected in 1984 under good plant-growth conditions. A reduction in nematode populations occurred in 1985 and 1986, possibly because of low soil-water conditions. There was a positive relationship between high soil water and maximum population densities of T. acutoides in the spring and fall of 1984, and between low soil water and minimum population densities of the nematode in 1985 and 1986. Pratylenchus neglectus populations were affected by soil water, although to a lesser degree than the ectoparasitic nematodes. Population densities of the three nematode species were significantly lower in the drier years of 1985 and 1986 than in 1984. Nematode populations were greater at the lower soil depths in the fall than in the spring or summer.     

Seasonal Changes in the Distribution of Photo-assimilated C in Young Pine Plants

Pinus strobus L. plants in their third year of growth were permitted to photoassimilate ¹⁴CO₂ for about 1 hour at monthly intervals between April and October, and the subsequent distribution of ¹⁴C in these plants was determined 8 hours, 1 month, 2 months or 4 months after photo-assimilation. In this way, the fate of ¹⁴CO₂ photo-assimilated during different months of the growing season was observed.

In the spring, old needles played a significant role in photo-assimilating ¹⁴CO₂ and exporting current photosynthate to the developing new shoots and roots. By July, the new shoot had replaced the old shoot both as the primary photo-assimilating part of the plant and as an exporter, particularly to the root.

The root received current photosynthate from the shoot throughout the entire growing season, although plant analysis only 8 hours after photo-assimilation did not always reveal this. Translocation of recent photosynthate from shoot to root was particularly high in August, September, and October.

The amounts of photo-assimilated ¹⁴C lost from the plants over a 4 month interval, principally through respiration and photorespiration, were about one-half of that absorbed during photo-assimilation, with the greatest loss occurring within the first month.

     

Distribution of ecosystem C and N within contrasting vegetation types in a semiarid rangeland in the Great Basin,USA

Semiarid sagebrush ecosystems are being transformed by wildfire, rangeland improvement techniques, and exotic plant invasions, but the effects on ecosystem C and N dynamics are poorly understood. We compared ecosystem C and N pools to 1 m depth among historically grazed Wyoming big sagebrush, introduced perennial crested wheatgrass, and invasive annual cheatgrass communities, to examine whether the quantity and quality of plant inputs to soil differs among vegetation types. Natural abundance δ¹⁵N isotope ratios were used to examine differences in ecosystem N balance. Sagebrush-dominated sites had greater C and N storage in plant biomass compared to perennial or annual grass systems, but this was predominantly due to woody biomass accumulation. Plant C and N inputs to soil were greatest for cheatgrass compared to sagebrush and crested wheatgrass systems, largely because of slower root turnover in perennial plants. The organic matter quality of roots and leaf litter (as C:N ratios) was similar among vegetation types, but lignin:N ratios were greater for sagebrush than grasses. While cheatgrass invasion has been predicted to result in net C loss and ecosystem degradation, we observed that surface soil organic C and N pools were greater in cheatgrass and crested wheatgrass than sagebrush-dominated sites. Greater biomass turnover in cheatgrass and crested wheatgrass versus sagebrush stands may result in faster rates of soil C and N cycling, with redistribution of actively cycled N towards the soil surface. Plant biomass and surface soil δ¹⁵N ratios were enriched in cheatgrass and crested wheatgrass relative to sagebrush-dominated sites. Source pools of plant available N could become ¹⁵N enriched if faster soil N cycling rates lead to greater N trace gas losses. In the absence of wildfire, if cheatgrass invasion does lead to degradation of ecosystem function, this may be due to faster nutrient cycling and greater nutrient losses, rather than reduced organic matter inputs.     

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     

Use of positron-emitting tracer imaging system for measuring the effect of salinity on temporal and spatial distribution of C tracer and coupling between source and sink organs

Salinity stress affects photosynthate partitioning between sources and sinks of plants, but how it affects these systems is less well understood. Because sources and sinks are closely tied, any adverse effect under suboptimal conditions on one of these is often misinterpreted for an effect on the other. Carbon partitioning is indispensable for stress resistance and good plant growth. In the present study, carbon partitioning in tomato plants (Lycopersicon esculentum L. cv. Momotarou) in a saline (NaCl) environment was studied by feeding radioactive ¹¹C and stable ¹³C isotopes. Pulse-chases were conducted to measure the spatial and temporal distribution of ¹³C. ¹³C was measured by a standard conventional technique, but ¹¹C distribution was monitored using a positron-emitting tracer imaging system (PETIS). Salt stress resulted in reduced carbon translocation toward roots. The majority of the photosynthate accumulated in the leaf. We also observed that the reduction in translocation of carbon occurred well before the salt stress symptoms of reduced photosynthesis and reduced plant growth in salt-exposed plants. The effect on sink activity was also shown by a decrease in stem diameter. In addition, PETIS analysis of ¹¹C translocation indicated that carbon translocation to roots was inhibited under salt conditions without a direct effect on leaf Na accumulation or osmotic stress. These results suggest that NaCl has direct effects on plants, inhibiting carbon partitioning within a few hours of salt exposure without inhibition of source activity.     

Effect of Single and Interplantings on Pathogenicity of Pratyenchus penetrans and P. neglectus to Alfalfa and Crested Wheatgrass

Alfalfa is a host of Pratylenchus penetrans and P. neglectus, whereas crested wheatgrass is a host of P. neglectus but not of P. penetrans. In a 120-day greenhouse experiment at 24 ñ 3 C, P. neglectus inhibited the growth of ''Lahontan'' alfalfa and ''Fairway'' crested wheatgrass. There were no differences in persistence and plant growth of alfalfa and crested wheatgrass, or reproduction of P. neglectus, in single plantings of alfalfa (AO) or crested wheatgrass (CWO), or in interplanted alfalfa and crested wheatgrass (ACW) treatments. On alfalfa, P. penetrans inhibited growth and reproduced more than did P. neglectus. Inhibition of plant growth and reproduction of P. penetrans was greater on alfalfa in AO than in ACW treatments. Pratylenchus penetrans did not reproduce on crested wheatgrass, but inhibited growth of crested wheatgrass in interplanted treatments and was avirulent in single planted treatments. Results were similar in a controlled growth chamber experiment at 15, 20, 25, and 30 C. Both nematode species inhibited alfalfa growth at all temperatures, and P. penetrans was more virulent than was P. neglectus to alfalfa at all temperatures and treatments. Plant growth inhibition and reproduction of P. penetrans on alfalfa in single and interplanted treatments were similar at 15-20 C, but were greater in single than in interplanted treatments at 25-30 C. Pratylenchus penetrans was avirulent to crested wheatgrass in the single planted treatments at all temperatures, but inhibited growth of crested wheatgrass in interplanted treatments at 20-30 C. Plant growth and reproduction of P. neglectus on crested wheatgrass was similar in single and interplanted treatments at 20-30 C and 15-30 C, respectively.     

Tillering in Tussock Grasses in Relation to Defoliation and Apical Bud Removal

Experiments with five caespitose grass species from temperateand tropical environments showed that the number of lateralshoots (tillers) which emerged following defoliation was notincreased by leaving a greater residual leaf area. Increasedavailability of photosynthate (and perhaps other resources)was effective, however, in increasing the rate of growth anddegree of flowering of new lateral shoots in one tropical species,Panicum maximum. In two temperate Agropyron tussock grasses, decapitation (apicalbud removal) did not stimulate lateral shoot growth. This indicatedthat apical dominance was not a factor preventing growth oflateral buds just prior to inflorescence emergence on the parenttillers. However, defoliation, where both terminal buds andfoliage were removed from the parent tillers stimulated lateralbud growth. Hormones other than those produced by the apicalbud or light quality or intensity may control lateral bud growthin these species. In contrast to the temperate species, lateralbud growth was stimulated by both decapitation and defoliationin the three tropical species. This response is consistent withthe model of correlative inhibition by apical dominance. Agropyron desertorum, Agropyron spicatum, Heteropogon contortus, Panicum maximum, Themeda triandra, crested wheatgrass, bluebunch wheatgrass, black speargrass, green panic grass kangaroo grass, apical dominance, tillering, regrowth, grazing, tussock grasses     

Effects of shading and of seasonal differences in weather on the growth, sugar content and sugar yield of sugar-beet crops

In 1967 and 1968 plots of sugar beet in two identically grown crops were shaded for successive four week periods starting on 13 June, 18 July and 22 August, and the growth of the plants compared with that on unshaded plots. At the beginning of each period in 1967 some shaded and unshaded plants received additional nitrogen, and in 1968 plants continuously shaded from June to September were compared with unshaded plants irrigated to equalize their water losses with those of the shaded plants. The weather in 1967 was sunny and dry and that in 1968 dull and wet, but the yields of dry matter, and particularly of sugar, of the unshaded crop in the 2 years were similar because, although net assimilation rate (E) was greater in 1967 than 1968, mean leaf area index (L) in 1968 was almost double that in the 1967 crop. Shading decreased the incoming radiation by 56%; it decreased E proportionally in 1967, increasing L slightly, but it decreased both E (by 44%) and L in 1968. The weights of dry matter and sugar in the roots of the shaded crop were consistently smaller at the end of shading and at final harvest in October in both years, and their weights, but not those of the tops or the whole plant, at final harvest over all shading treatments in both years were proportional to the amount of radiation received by the crop between June and September. Although shading greatly decreased the supply of photosynthate to the roots, it did not change the sugar content percentage of dry matter, except in the early stages of growth when the sugar content was rapidly increasing. Sugar content percentage of fresh weight of the roots was consistently decreased by shading, wholly because water content was increased relative to dry matter. Therefore the sugar yield of shaded plants was less because the dry weights of the roots were less, not because the partition of photosynthate between sugar storage and root growth changed. There appears to be a mechanism within the root operating over a wide range of photosynthate supply that maintains a nearly constant proportion of sucrose to non-sugar, contrary to the hypothesis that sugar stored in the roots is photosynthate in excess of what can be used in growth of the plant. However, in the extreme condition of continuous shading which drastically decreased the dry weights of all parts of the plant, sugar percentage of dry matter in the roots was decreased, but only from 80 to 70%. In 1967 extra nitrogen applied at the start of shading increased L and the dry weight of the tops in all periods, but had no effect on the dry weight of roots. Because 1968 was a wet year the irrigation treatment had no effect on E or L except for a slight increase in L during the first period; it had no effect on plant dry weight. Both irrigation and additional nitrogen decreased the sugar content percentage of fresh weight of the roots only by altering their water content; sugar percentage of dry matter was unaffected. None of these changes persisted until the final harvest in October.     

Responses of tree fine roots to temperature

Soil temperature can influence the functioning of roots in many ways. If soil moisture and nutrient availability are adequate, rates of root length extension and root mortality increase with increasing soil temperature, at least up to an optimal temperature for root growth, which seems to vary among taxa. Root growth and root mortality are highly seasonal in perennial plants, with a flush of growth in spring and significant mortality in the fall. At present we do not understand whether root growth phenology responds to the same temperature cues that are known to control shoot growth. We also do not understand whether the flush of root growth in the spring depends on the utilization of stored nonstructural carbohydrates, or if it is fueled by current photosynthate. Root respiration increases exponentially with temperature, but Q ₁₀ values range widely from c . 1.5 to > 3.0. Significant questions yet to be resolved are: whether rates of root respiration acclimate to soil temperature, and what mechanisms control acclimation if it occurs. Limited data suggest that fine roots depend heavily on the import of new carbon (C) from the canopy during the growing season. We hypothesize that root growth and root respiration are tightly linked to whole-canopy assimilation through complex source–sink relationships within the plant. Our understanding of how the whole plant responds to dynamic changes in soil temperature, moisture and nutrient availability is poor, even though it is well known that multiple growth-limiting resources change simultaneously through time during a typical growing season. We review the interactions between soil temperature and other growth-limiting factors to illustrate how simple generalizations about temperature and root functioning can be misleading.     

Partitioning of shoot and root dry matter and carbohydrates in bean plants suffering from phosphorus, potassium and magnesium deficiency

The influence of varied supply of phosphorus (10 and 250 mmolP m^–3) potassium (50 and 2010 mmol K m^–3) and magnesium(20 and 1000 mmol Mg m^–3) on the partitioning of dry matterand carbohydrates (reducing sugars, sucrose and starch) betweenshoots and roots was studied in bean (Phaseolus vulgaris) plantsgrown in nutrient solution over a 12 d period. Shoot and rootgrowth were quite differently affected by low supply of P, K,and Mg. The shoot/root dry weight ratios were 4.9 in the control(sufficient plants), 1.8 in P-deficient, 6.9 in K-deficientand 10.2 in Mg-deficient plants. In primary (source) leaves,but not in trifoliate leaves, concentrations of reducing sugars,sucrose and starch were also differently affected by low nutrientsupply. In primary leaves under K deficiency and, particularlyMg deficiency, the concentrations of sucrose and reducing sugarswere much higher than in control and P-deficient plants. Magnesiumdeficiency also distinctly increased the starch concentrationin the primary leaves. In contrast, in roots, the lowest concenfrationsof sucrose, reducing sugars and starch were found in Mg-deficientplants, whereas the concentrations of sucrose and starch wereparticularly high in P-deficient plants. There was a close relationshipbetween shoot/root dry weight ratios and relative distributionof total carbohydrates (sugars and starch) in shoot and roots.Of the total amounts of carbohyd rates per plant, the followingproportions were parti tioned to the roots: 22.7% in P-deficient,15.7% in control, 3.4% in K-deficient and 0.8% in Mg-deficientplants. The results indicate a distinct role of Mg and K in the exportof photosynthates from leaves to roots and suggest that alterationin photosynthate partitioning plays a major role in the differencesin dry matter distribution between shoots and roots of plantssuffering from mineral nutrient deficiency. Key words: Bean, carbohydrates, magnesium nutrition, phosphorus nutrition, potassium nutrition, shoot/root growth     

Rhizome dynamics and resource storage in Phragmites australis

Seasonal changes in rhizome concentrations of total nonstructural carbohydrates (TNC), water soluble carbohydrates (WSC), and mineral nutrients (N, P and K) were monitored in two Phragmites australis stands in southern Sweden. Rhizome biomass, rhizome length per unit ground area, and specific weight (weight/ length ratio) of the rhizomes were monitored in one of the stands.Rhizome biomass decreased during spring, increased during summer and decreased during winter. However, changes in spring and summer were small (< 500 g DW m-2) compared to the mean rhizome biomass (approximately 3000 g DW m^–2). Winter losses were larger, approximately 1000 g DW m-2, and to a substantial extent involved structural biomass, indicating rhizome mortality. Seasonal changes in rhizome length per unit ground area revealed a rhizome mortality of about 30% during the winter period, and also indicated that an intensive period of formation of new rhizomes occurred in June.Rhizome concentrations of TNC and WSC decreased during the spring, when carbohydrates were translocated to support shoot growth. However, rhizome standing stock of TNC remained large (> 1000 g m^–2). Concentrations and standing stocks of mineral nutrients decreased during spring/ early summer and increased during summer/ fall. Only N, however, showed a pattern consistent with a spring depletion caused by translocation to shoots. This pattern indicates sufficient root uptake of P and K to support spring growth, and supports other evidence that N is generally the limiting mineral nutrient for Phragmites.The biomass data, as well as increased rhizome specific weight and TNC concentrations, clearly suggests that reloading of rhizomes with energy reserves starts in June, not towards the end of the growing season as has been suggested previously. This resource allocation strategy of Phragmites has consequences for vegetation management.Our data indicate that carbohydrate reserves are much larger than needed to support spring growth. We propose that large stores are needed to ensure establishment of spring shoots when deep water or stochastic environmental events, such as high rhizome mortality in winter or loss of spring shoots due to late season frost, increase the demand for reserves.     

Off-season uptake of nitrogen in temperate heath vegetation

In this field study we show that temperate coastal heath vegetation has a significant off-season uptake potential for nitrogen, both in the form of ammonium and as glycine, throughout winter. We injected ¹⁵N-ammonium and ¹⁵N 2×(¹³C)-glycine into the soil twice during winter and once at spring. The winter temperatures were similar to those of an average winter in the northern temperate region of Europe, with only few days of soil temperatures below zero or above 5°C. The vegetation, consisting of the evergreen dwarf shrub Calluna vulgaris, the deciduous dwarf shrub Salix arenaria, and the graminoids Carex arenaria and Deschampsia flexuosa, showed high root uptake of both forms of nitrogen, both 1 day after labelling and after a month, in species specific temporal patterns. Plant uptake of ¹³C was not significant, providing no further evidence of intact uptake of glycine. Translocation of the labelled nitrogen to shoots was generally evident after 1 month and increased as spring approached, with different translocation strategies in the three plant functional types. Furthermore, only the graminoids showed shoot growth during winter. Increasing plant nitrogen concentration from fall to spring at temperate heaths may, hence, be due to nitrogen uptake. Our results suggest that the potential for nitrogen uptake in plants at winter is of the same order of magnitude as at summer. Hence, winter nitrogen uptake in ecosystems in the temperate/boreal region should be considered when making annual nitrogen budgets of heath ecosystems, and the view of plant nutrient uptake as low in this climatic region during winter should be revised.     

Dynamics of growth,carbon and nutrient translocation in Zizania latifolia

We studied the seasonal resource dynamics between organs of wild rice (Zizania latifolia (Griseb.) Turcz. ex Stapf.) to obtain a better understanding of its growth dynamics, carbon and nutrient translocation. The results of observation from January 2002 to February 2004 showed the shoot density markedly increased after emergence of shoots at the end of March until May (up to 800 ind/m²). However the shoot mortality due to self-thinning reduced the total new shoots by more than 70% by the end of July. Thereafter, the shoot density was nearly constant with the aboveground biomass peaking at the end of August. In the late winter, the rhizome biomass declined by respiration loss to about 25% of its peak value. Meanwhile the decline in rhizome reserves from January to the end of April was about 20%. This small reduction compared with other perennial emergent species implies that there is a lower contribution of rhizome reserves to support new shoot formation. The initial heterotrophic growth of new shoots based on the rhizome resources lasted for a short period, then switched to autotrophic growth at the end of April or the beginning of May. Thus, in most periods of foliage development, nutrients were obtained mostly from soil through uptake by roots, not through resource allocation of the rhizome. In autumn, the standing dead shoots retained most of the nutrients and carbohydrates without translocating downwards. This suggests that in practice, the plant can remove nutrients from sediment more efficiently than other emergent plants.     

Phytoremediation of Pentachlorophenol in the Crested Wheatgrass (Agropyron cristatum × desertorum) Rhizosphere

Pentachlorophenol (PCP) is a widespread, highly toxic contaminant of soil and water that is generally recalcitrant to microbial breakdown and thus may be considered a good candidate for phytoremediation. PCP toxicity and rates of mineralization were compared in crested wheatgrass seedlings that were either sterile or root-inoculated with microbial consortia derived from soil at a PCP-contaminated site. Inoculated seedlings were more tolerant to PCP and mineralized threefold more ¹⁴C-PCP than sterile seedlings. Only 10% of the recovered radioactivity from sterile seedlings represented mineralized PCP, indicating that rhizosphere microorganisms are primarily responsible for PCP mineralization. The levels of PCP degradation exhibited by several microbial consortia and isolates in liquid culture were not correlated with their ability to protect crested wheatgrass seedlings from PCP toxicity. Most probable number estimates showed that the presence of crested wheatgrass root exudates enhanced the number of PCP-degrading microorganisms by 100-fold in liquid culture, indicating that exudate components provide some nutritive benefit, possibly as PCP co-metabolites. A close association of plants and rhizosphere microorganisms appears to be necessary for crested wheatgrass survival in PCP-contaminated soil, although understanding the molecular details of this association requires further research.