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.     

Effect of defoliation intensity on aboveground and belowground relative growth rates

According to a simple growth model, grazed and ungrazed plants may have equal absolute growth rates provided that the relative growth rate (RGR) of grazed plants increases exponentially with grazing intensity (proportion of biomass removed). This paper reports results from an experiment designed to determine whether plants of two grass species subjected to a gradient of defoliation intensities, from 0 to 100% aboveground biomass removal, showed such a response. The relationship between aboveground RGR and defoliation intensity was exponential and closely matched the theoretical relationship of equal absolute growth rate. Thus, plants showed the same aboveground growth regardless of defoliation intensity thanks to an exponential stimulation of RGR by defoliation. Belowground RGR was depressed by defoliation of more than 20% of the above-ground biomass. In spite of the drastic modification imposed by the treatments on the relative proportions of different plant parts, after a 42-day recovery period basic allometric relationships, such as root:shoot and leafarea: weight ratios, were not affected by defoliation intensity. Exponential aboveground compensatory responses represent a key feedback process resulting in constant aboveground growth regardless of defoliation intensity and appear to be a simple consequence of strong commitments to certain allometric relationships.     

The effects of the spatial pattern of defoliation on regrowth of a tussock grass

Summary The spatial pattern of foliage removal from a tussock grass can influence regrowth through effects on daily carbon gain (CER_d). This field study examined the extent to which tussock photosynthetic responses to different defoliation patterns were associated with changes in whole-canopy attributes (e.g., foliage age structure, canopy light microclimate). During the spring growing season, 60% of the green foliage area was removed from individual Agropyron desertorum tussocks with scissors in different spatial patterns. These patterns represented extremes of defoliation patterns that might be inflicted by natural herbivores. Tussock photosynthesis (per unit foliage area) at high light (2000 mol photons m^–2 s^–1 between 400 and 700 nm; P₂₀₀₀) increased following clipping with all defoliation patterns. The increases in P₂₀₀₀ were greater when leaves were removed from low in the tussock (older leaves) than if leaves high in the canopy (younger leaves) were removed. These relative changes of P₂₀₀₀ among clipping patterns paralleled the responses of CER_d and regrowth from an earlier study. Furthermore, the changes in P₂₀₀₀ corresponded with increases in the proportion of foliage within the tussocks that was directly illuminated at midday. The greater photosynthesis of tussocks after lower-leaf removal was directly related to a higher proportion of younger foliage and a smaller fraction of foliage shaded within the tussock. In a dense canopy, such as these grass tussocks, the influence of defoliation on whole-canopy attributes may be of primary importance to whole-plant photosynthetic responses.     

Ecological conditions that determine when grazing stimulates grass production

Summary We report the results of a pot experiment that examined the effects of three ecologically important factors controlling plant growth rates in savanna grasslands: defoliation, soil nitrogen and soil water availability. The experiment was conducted in the Amboseli region in east Africa, and was designed to simulate natural conditions as far as possible, using local soils and a grass species that is heavily grazed by abundant large herbivores. Productivity by different plant components was reduced, stimulated or unchanged by defoliation, depending on specific watering and fertilization treatments. Total above-ground production was stimulated by defoliation and was maximized at moderate clipping intensities, but this was statistically significant only when plants were watered infrequently (every 8 days), and most important, periods between clipping events were extended (at least 24 days). Under these conditions, plant growth rates were limited by water availability at the time of clipping, and soil water conserved in clipped, compared to unclipped plants. Within a given fertilization treatment, whole-plant production was never stimulated by defoliation because root growth was unaffected or inhibited by clipping. However, when fertilization was coupled to defoliation, as they are in the field, whole-plant production by fertilized and moderately clipped plants exceeded production by infertilized, unclipped plants. Under this interpretation, maximum whole-plant production coincided with optimum conditions for herbivores (maximum nitrogen concentration in grass leaves) when watering was frequent, and plants were moderately defoliated. However, these conditions were not the same as those that maximized relative above-ground stimulation of growth (infrequent watering and clipping).The results indicate that above-ground grass production can be stimulated by grazing, and when that is likely to occur. However, the results emphasize that plant production responses to defoliation can vary widely, contigent upon a complex interaction of ecological factors.     

The effect of shading on photosynthesis,growth, and regrowth following defoliation for Bromus tectorum

Summary The effect of full sunlight, 60%, or 90% attenuated light on photosynthetic rate, growth, leaf morphology, dry weight allocation patterns, phenology, and tolerance to clipping was examined in the glasshouse for steppe populations of the introduced grass, Bromus tectorum. The net photosynthetic response to light for plants grown in shade was comparable to responses for plants grown in full sunlight. Plants grown in full sunlight produced more biomass, tillers and leaves, and allocated a larger proportion of their total production to roots than plants grown in shade. The accumulation of root and shoot biomass over the first two months of seedling growth was primarily responsible for the larger size at harvest of plants grown in full sunlight. Plants grown under 60% and 90% shade flowered an average of 2 and 6 weeks later, respectively, than plants grown in full sunlight. Regrowth after clipping was greater for plants grown in full sunlight compared to those grown in shade. Even a one-time clipping delayed flowering and seed maturation; the older the individual when leaf area was removed, the greater the delay in its phenology. Repeated removal of leaf area was more frequently fatal for plants in shade than in full sunlight. For plants originally grown in full sunlight, regrowth in the dark was greater than for shaded plants and was more closely correlated to non-flowering tiller number than to plant size. This correlation suggests that etiolated regrowth is more likely regulated by the number of functional meristems than by differences in the size of carbohydrate pools. Thus, shading reduces the rate of growth, number of tillers, and ability to replace leaf area lost to herbivory for B. tectorum. These responses, in turn, intensify the effect of competition and defoliation for this grass in forests. B. tectorum is largely restricted to forest gaps at least in part because of its inability to acclimate photosynthetically, the influence of shade on resource allocation, and the role of herbivory in exacerbating these effects.     

Flooding effects on rapid responses of the invasive plant Alternanthera philoxeroides to defoliation

In experiments under controlled growth conditions it was examined how flooding affected the responses of the invasive plant Alternanthera philoxeroides to defoliation. In drained and flooded conditions, plants were subjected to five defoliation levels: 0, 10, 50, 90% removal of leaf tissue and apex removal (90% leaf tissue plus apical bud removal). Plants were harvested weekly for five weeks. In drained conditions, plant biomasses including total biomass, shoot biomass and root biomass after 50% defoliation rapidly recovered to the control plant level. They were significantly lower for the 90% defoliation and apex removal treatments compared to control plants throughout the experiment. In flooded conditions, total biomass and shoot biomass after 50% defoliation, 90% defoliation, and apex removal treatments could return to control plant levels before the end of the experiment. In 90% defoliation and apex removal treatments root to shoot biomass ratios of both drained and flooded plants were initially much higher than in control plants, but the difference disappeared rapidly. The final biomasses decreased with increased defoliation intensity in drained conditions, but no significant difference was generally found in any of the defoliation treatments in flooded conditions. The rapid re-growth of A. philoxeroides plants after defoliation may partly be responsible for its invasion success. However, defoliation capable of removing 90% of the leaf tissue may be desirable in restricting the growth of this invasive species in drained conditions.     

Toxic effects of Pb2+ on the growth and mineral nutrition of signal grass (Brachiaria decumbens) and Rhodes grass (Chloris gayana)

Although grasses are commonly used to revegetate sites contaminated with lead (Pb), little is known regarding the Pb-tolerance of many of these species. Using dilute solution culture to mimic the soil solution, the growth of signal grass (Brachiaria decumbens Stapf cv. Basilisk) and Rhodes grass (Chloris gayana Kunth cv. Pioneer) was related to the mean activity of Pb²⁺ {Pb²⁺} in solution. There was a 50% reduction in fresh mass of signal grass shoots at 5 μM {Pb²⁺} and at 3 μM {Pb²⁺} for the roots. Rhodes grass was considerably more sensitive to Pb in solution, with shoot and root fresh mass being reduced by 50% at 0.5 μM {Pb²⁺}. The higher tolerance of signal grass to Pb appeared to result from the internal detoxification of Pb, rather than from the exclusion of Pb from the root. At toxic {Pb²⁺}, an interveinal chlorosis developed in the shoots of signal grass (possibly a Pb-induced Mn deficiency), whilst in Rhodes grass, Pb²⁺ caused a bending of the root tips and the formation of a swelling immediately behind some of the root apices. Root hair growth did not appear to be reduced by Pb²⁺ in solution, being prolific at all {Pb²⁺} in both species.     

Mechanisms for the increase in phosphorus uptake of waterlogged plants: soil phosphorus availability, root morphology and uptake kinetics

Waterlogging frequently reduces plant biomass allocation to roots. This response may result in a variety of alterations in mineral nutrition, which range from a proportional lowering of whole-plant nutrient concentration as a result of unchanged uptake per unit of root biomass, to a maintenance of nutrient concentration by means of an increase in uptake per unit of root biomass. The first objective of this paper was to test these two alternative hypothetical responses. In a pot experiment, we evaluated how plant P concentration of Paspalum dilatatum, (a waterlogging-tolerant grass from the Flooding Pampa, Argentina) was affected by waterlogging and P supply and how this related to changes in root-shoot ratio. Under both soil P levels waterlogging reduced root-shoot ratios, but did not reduce P concentration. Thus, uptake of P per unit of root biomass increased under waterlogging. Our second objective was to test three non-exclusive hypotheses about potential mechanisms for this increase in P uptake. We hypothesized that the greater P uptake per unit of root biomass was a consequence of: (1) an increase in soil P availability induced by waterlogging; (2) a change in root morphology, and/or (3) an increase in the intrinsic uptake capacity of each unit of root biomass. To test these hypotheses we evaluated (1) changes in P availability induced by waterlogging; (2) specific root length of waterlogged and control plants, and (3) P uptake kinetics in excised roots from waterlogged and control plants. The results supported the three hypotheses. Soil P avail-ability was higher during waterlogging periods, roots of waterlogged plants showed a morphology more favorable to nutrient uptake (finer roots) and these roots showed a higher physiological capacity to absorb P. The results suggest that both soil and plant mechanisms contributed to compensate, in terms of P nutrition, for the reduction in allocation to root growth. The rapid transformation of the P uptake system is likely an advantage for plants inhabiting frequently flooded environments with low P fertility, like the Flooding Pampa. This advantage would be one of the reasons for the increased relative abundance of P. dilatatum in the community after waterlogging periods. Received: 15 February 1997 / Accepted: 20 May 1997     

Phosphorus availability and rootstock affect copper-induced damage to the root ultra-structure of Citrus

The control of several citrus diseases requires continuous applications of fungicides containing copper (Cu) which favor to the accumulation of this metal in the soil. Therefore, the evaluation of how nutrient availability and rootstock interact with Cu toxicity in the citrus trees is required to maintain sustainability of fruit production in Cu-contaminated soils. Valencia orange trees on Sunki mandarin (SM) or Swingle citrumelo (SC) rootstock were grown in nutrient solutions combining adequate Cu (1.0 μmol L⁻¹), excess Cu (50.0 μmol L⁻¹), deficient phosphorus (P) (0.01 mmol L⁻¹) and sufficient P (0.5 mmol L⁻¹). The excess Cu reduced root and shoot growth, chlorophyll and relative water content in the leaves of the trees compared to those under adequate Cu supply. Furthermore, excess Cu caused severe damage to the root ultra-structure, characterized by the degeneration of the middle lamella and the presence of a thin and sinuous cell wall, as well as, starch accumulation in the plastids, disruption of the mitochondrial membranes and cellular plasmolysis. The damage caused by excess Cu in the cell wall and middle lamella on the root cells of SC was less severe than SM. Sufficient P supply improved the structure of the cell wall and middle lamella of trees subjected to excess Cu in comparison to P-deficient ones. Thus, the occurrence of more preserved cell wall and middle lamella supports the idea that sufficient P availability in the rooting medium and the use of SC rootstock might contribute to increase the ability of young citrus trees to cope with Cu toxicity.     

Compensatory growth responses to clipping defoliation in Leymus chinensis (Poaceae) under nutrient addition and water deficiency conditions

Compensatory growth responses of Leymus chinensis, a dominant species in Inner Mongolia steppe, to clipping defoliation were evaluated in a pot-cultivated experiment under different nutrient (N and P) and water availability conditions. Leymus chinensis exhibited over-compensatory growth at the light and moderate clipping intensities (20% and 40% aerial mass removed) with a greater accumulated aboveground biomass, higher relative growth rate (RGR), more rhizomatic tillers and a stimulation of compensatory photosynthesis to the remnant leaves as compared with those of the unclipped plants. Intense clipping (80% aerial mass removed), which removed most of the aboveground tissues, greatly reduced the growth of aboveground biomass in comparison with that of the unclipped plants. Nitrogen addition only slightly improved the biomass production and RGR in light and moderately clipped plants, and it did not allow plants in the intense clipping condition to over-compensate. Phosphorus addition had no obvious influences on the growth and physiological responses to clipping defoliation. These results indicated that nutrient addition could not compensate for the negative effects of severe clipping on the defoliated grass. On the other hand, there were no distinct positive responses under water deficiency condition for L. chinensis at all clipping intensities with a significant reduction of aboveground and belowground biomass, lower RGR, fewer rhizomatic tillers, and a lower net photosynthetic rate than other wet treatments. Additionally, the chlorophyll contents of remnant leaves gradually increased with the increase of clipping intensities in each treatment. In conclusion, although L. chinensis could compensate for tissues removal by some morphological and physiological responses, intense clipping and drought can result in a significant decrease of biomass and growth rate, even under enriched nutrition conditions.     

水产动物继饥饿或营养不足后的补偿生长研究进展

综述了水产动物继饥饿或营养不足后的补偿生长研究进展,其中包括补偿生长的程度、影响因素、生理学机制、补偿生长研究的实验设计、补偿生长过程中生长率和生化组成的变化及存在的问题和应用前景。     

淡水养殖太平洋鲑循环饥饿后补偿性生长效果研究

用16.1%脂肪,38.1%蛋白质含量日粮饲养108尾初始重约为240g的太平洋鲑(Oncorhynchusspp.)于0.25m3的水族箱中64d,水温为15.5±3.7℃。实验分6组,分别为对照组(每天投喂),实验1组(隔天投喂),实验2组(隔2天投喂2天),实验3组(隔4天投喂4天),实验4组(隔8天投喂8天),实验5组(隔16天投喂16天)。每组设3个平行水族箱,每箱6尾鱼。研究淡水养殖太平洋鲑多重周期饥饿后补偿性生长效果。实验结果表明:(1)各试验组太平洋鲑成活率均为100%。实验1、2、3组太平洋鲑鱼体增重接近对照组,其恢复摄食期间特定生长率、摄食率、食物转化率均显著或极显著高于对照组(P<0.05或0.01)。而实验4、5组鱼恢复摄食期间虽摄食率极显著高于对照组(P<0.01),但其鱼体增重、特定生长率、食物转化率均极显著低于对照组(P<0.01);(2)实验各组鱼肥满度、肝体比、肝脏脂肪和糖原含量、肌肉中脂肪含量较对照组均有不同程度下降,肝脏脂肪中总饱和脂肪酸比例上升,而总多不饱和脂肪酸比例下降;(3)实验1、2、3组血浆中甘油三酯、胆固醇和低密度脂蛋白显著低于对照组,而葡萄糖、血清中甲状腺激素T4浓度显著高于对照组(P<0.05)。实验结果表明,初重约240g太平洋鲑饥饿1—4d,再循环投喂相同时间64d后,获得了接近完全补偿生长效果,表现为其恢复摄食期间摄食率和食物转化率明显上升,生长速率明显加快,饲料报酬明显提高,鱼体增重接近持续喂食的对照组,养殖效益明显提高。但饥饿8—16d再循环投喂相同时间后,表现为无补偿生长效应,食物转化率和生长速率明显下降,鱼体增重极显著低于持续喂鱼的对照组。     

Clipping defoliation and nitrogen addition shift competition between a C3 grass (Leymus chinensis) and a C4 grass (Hemarthria altissima)

• Human‐induced disturbances, including grazing and clipping, that cause defoliation are common in natural grasslands. Plant functional type differences in the ability to compensate for this tissue loss may influence interspecific competition.
• To explore the effects of different intensities of clipping and nitrogen (N) addition on compensatory growth and interspecific competition, we measured accumulated aboveground biomass (AGB), belowground biomass (BGB), tiller number, non‐structural carbohydrates concentrations and leaf gas exchange parameters in two locally co‐occurring species (the C₃ grass Leymus chinensis and the C₄ grass Hemarthria altissima) growing in monoculture and in mixture.
• For both grasses, the clipping treatment had significant impacts on the accumulated AGB, and the 40% clipping treatment had the largest effect. BGB gradually decreased with increasing defoliation intensity. Severe defoliation caused a significant increase in tiller number. Stored carbohydrates in the belowground biomass were mobilised and transported aboveground for the growth of new leaves to compensate for clipping‐induced injury. The net CO₂ assimilation rate (A) of the remaining leaves increased with clipping intensity and peaked under clipping intensities of 20% or 40%. Nitrogen addition, at a rate of 10 g·N·m^?2·year^?1, enhanced A of the remaining leaves and non‐structural carbohydrate concentrations, which benefited plant compensatory growth, especially for the C₃ grass. Under the mixed planting conditions, the clipping and N addition treatments lowered the competitive advantage of the C₄ grass.
• The results suggest that a combination of defoliation and N deposition have the potential to benefit the coexistence of C₃ and C₄ grasses.

     

Trade-off between cadmium tolerance and relative growth rate in 10 grass species

Plant species exhibit great differences in heavy metal accumulation and tolerance. In this study we compared interspecific differences in responses to Cadmium (Cd) stress among 10 C₃ grass species by growing hydroponically under the conditions of different Cd treatments (0, 5, 10 and 50 μM). Responses of plant shoot dry mass to Cd treatments (resistance) were separated into avoidance (the response of shoot Cd concentration to Cd treatments) and tolerance (the response of shoot dry mass to shoot Cd concentration). The relative growth rate (RGR) and leaf structural properties of plants were measured under optimum growth condition. There were large differences in resistance, avoidance and tolerance among the species. Avoidance and tolerance were attributed independently to total Cd resistance. Resistance and tolerance were correlated negatively with RGR, leaf water content (LW), specific leaf area (SLA), leaf elongation rate (LER), and leaf length (LL), but the leaf dry matter concentration (LMDC), and nuclear DNA content showed a positive correlation with resistance and tolerance. These results indicate a trade-off between growth rate and tolerance to Cd stress. Species with higher Ca/Mg ratios showed low avoidance (r = −0.943***), suggesting uptake inhibition by Ca²⁺ to develop avoidance against Cd stress.     

Characteristic responses of three tropical legumes to the inoculation of two species of VAM fungi in Andosol soils with different fertilities

The growth and mineral nutrition responses were evaluated of three tropical legumes, cowpea (Vigna unguiculata L. cv Kuromame), pigeonpea [Cajanus cajan L. (Millsp.) cv ICPL 86009] and groundnut (Arachis hypogaea cv Nakateyutaka) inoculated with two different species of VAM fungi, Glomus sp. (Glomus etunicatum-like species) and Gigaspora margarita, and grown in Andosols with different fertilities [Bray II-P: topsoil (72 ppm), subsoil (<0.1 ppm)]. Percent fungal root colonization was high in cowpea and groundnut but relatively low in pigeonpea in both soil types. Despite the low rate of root infection, significant growth responses were produced, especially in the inoculated pigeonpea plant. In all legumes, shoot dry matter production was favoured by the inoculations. Increases in shoot biomass due to mycorrhizae were greater in the subsoil than in the topsoil. Mycorrhization raised shoot concentrations of P and Ca (in cowpea and groundnut) and P and K (in pigeonpea) in the topsoil. Whereas the P concentration in shoots in the subsoil was not positively affected by VAM fungi, particularly in cowpea and pigeonpea, the concentration of K in such plants was significantly increased by VAM treatment. The results also showed that mycorrhizal enhancement of shoot micronutrient concentrations was very rare in all plants, with negative effects observed in certain cases. Cu concentration, in particular, was not affected by VAM formation in any of the plants, and Mn and Fe in pigeonpea and groundnut, respectively, remained the same whether plants were mycorrhizal or not. In both soils the three legumes responded to Glomus sp. better than to Gigaspora margarita, and the effects of the VAM fungi on each of the crops relative to the controls were greater in the subsoil than in the topsoil. However, shoot growth of groundnut was not affected as much as cowpea and pigeonpea by the type of soil used. In spite of the relatively low infection of its root, pigeonpea was generally the most responsive of the three legume species in terms of mycorrhizal/nonmycorrhizal ratios.     

Epidermal growth factor promotes oligodendrocyte process formation and regrowth after injury

Oligodendrocytes (OLs) form myelin within the central nervous system and are targets in numerous demyelinating diseases and injuries. OLs grown in culture maintain the developmental timetable which occurs in vivo and mature into cells with a relatively normal phenotype. In this study, cultured cells are used to test whether EGF can modulate process formation in OLs both before and after transection injury. EGF had no effect on the formation of new processes by OLs at any stage of development. To test the effect of EGF on process outgrowth after injury, mature OLs were selected and injured by laser transection of a single process, then imaged at 24-h intervals for 120 h. EGF promoted the recovery and regrowth of injured processes and also significantly increased outgrowth in uninjured processes. As well, it increased the number of new sprouts formed by OLs after injury. Results suggest that the effects of EGF on process outgrowth are a consequence of EGF interaction with a signaling pathway that is specifically activated within injured OLs. The potent effect of EGF on OL process formation after an injury suggests that modulation of the signaling pathways involved might provide a mechanism to promote remyelination.