The role of roots and cotyledons as storage organs in early stages of establishment in Quercus crispula: a quantitative analysis of the nonstructural carbohydrate in cotyledons and roots

Quercus seedlings have hypogeal cotyledons and tap roots, both of which act as storage organs. The importance of the storage function in the two organs may change as the seedling develops. Therefore, changes in carbohydrate reserves in cotyledons and roots of Q. crispula grown under 40 % and 3 % of full light from shoot emergence to the completion of the first leaf flush were monitored. In addition, a shoot-clipping treatment was performed to examine the relative contribution of the cotyledons and tap roots to resprouting. Cotyledons maintained large amounts of nonstructural carbohydrates during shoot development, and carbohydrates were still present in the cotyledons during the final phase of leaf flush. In addition, a notable increase in the amount of carbohydrates was observed in tap roots before leaf flush at both light levels. Since root development occurred before leaf flush, even in plants grown under 3 % light, the carbohydrate found in them presumably originated from seed reserves and was translocated to roots as storage reserves. When shoots were clipped at the leaf flushing stage, the amount of carbohydrate decreased only in the cotyledons after resprouting, suggesting that cotyledons act as the main storage organs during shoot development stages. However, it could be advantageous as a 'risk avoidance strategy' for the seedlings to store reserves in both cotyledons and roots, since cotyledons may be removed by predators during shoot development.     

Resprouting ability of<Emphasis Type="Italic"> Quercus crispula</Emphasis> seedlings depends on the vegetation cover of their microhabitats

To examine the effects of vegetation cover on the resprouting abilities of Quercus crispula seedlings, in each of three consecutive years, we artificially clipped seedlings growing in microhabitats with differing degrees of vegetation cover. We also investigated the relationship between the level of total nonstructural carbohydrate (TNC) and resprouting ability. Seedlings with clipped shoots in gaps produced larger resprouting shoots than those in the understory. Moreover, both the percentage of resprouting seedlings and the survival ratio in seedlings with clipped shoots were negatively correlated with the degree of vegetation cover. Seedlings stored high levels of TNC, especially in their roots, and their TNC levels were negatively correlated with the degree of vegetation cover. There were also positive relationships between the TNC levels in their roots and the degree of resprouting. Hence, we conclude that release from vegetation cover enhanced the resprouting ability of Q. crispula seedlings by increasing their levels of stored carbohydrate. The key variables affected were the resprouting ratio (the proportion of seedlings capable of producing new shoots) and the size of the resprouted shoots.     

Effects of bicarbonate and high pH on growth of Zn-efficient and Zn-inefficient genotypes of rice,wheat and rye

We studied the source of the nitrogen used for the growth and resprouting of holm-oak (Quercus ilexL.), and the contribution of nitrogen and carbohydrate root reserves to these processes. Three-year-old plants were grown in a greenhouse with either a sufficient or restricted nitrogen supply for one year. Half the individuals were subjected to shoot excision to provoke resprouting, and a ¹⁵N solution was given to these plants and to controls for two months. Nitrogen, Total Non-structural Carbohydrate (TNC), Total Soluble Protein content, and ¹⁵N and ¹³C composition were determined, and histological analyses of woody tissue were performed. Our results show that N-deprived plants used nitrogen from root reserves to support a growth rate similar to that of non-deprived plants. However, deprived plants lost their resprouting capacity in spite of the high TNC accumulation and nitrogen resupply to the soil. After the supply of nitrogen was restored to N-deprived plants, this nutrient mainly accumulated in under-ground organs, which limited the above-ground growth. Resprouting plants first remobilized the nitrogen stored in roots, and thereafter took it up from the solution. The root-crown region did not behave as a specialised reserve organ in three-year-old Quercus ilex L. plants.     

Responses of <Emphasis Type="Italic">Caragana korshinskii</Emphasis> Kom. to shoot removal: mechanisms underlying regrowth

Caragana korshinskii Kom. a dominant member of desert flora in north-western China, is often subjected to aboveground shoot destruction but is very successful in its rapid recovery. We investigated the physiological basis for resprouting by comparing shoot elongation, leaf-nutrient content, pre-dawn leaf-water potential (LWP), root non-structural carbohydrate (TNC), and photosynthetic rate of first-year resprouts with those of adjacent undamaged individuals. C. korshinskii resprouts had a significantly higher rate of shoot elongation. Plant responses associated with enhanced shoot elongation included (1) improved water status, (2) drawing upon more TNC from roots to support aboveground shoot regrowth, (3) enhanced photosynthetic rate owing to improved water status and increased nutrient content in leaves, and (4) allocating more photosynthate to vegetative production without reproduction costs. Maintaining an active root system after shoot removal may be the foundation which engenders these mechanisms underlying rapid regrowth of C. korshinskii in the disturbed environment.     

Postfire regeneration of resprouting mountain fynbos shrubs: differentiating obligate resprouters and facultative seeders

Plant species in Mediterranean-type climate regions have a diversity of traits that facilitate their persistence under a given fire regime. Obligate resprouters (OR) are dependent on resprouting to persist through a burn episode, as their seeds are killed by fire. Facultative seeders (FS) combine strategies by resprouting and recruiting new seedlings after fire. We hypothesised that these life history differences would lead to differential resprout success and we predicted that OR would be more successful than FS. We performed a 2-year study to assess resprout success of co-occurring Western Cape mountain fynbos FS and OR species, and to determine predictors of resprout success following a wildfire. All the FS species recruited seedlings postfire, whereas the OR did not. OR demonstrated near complete (99 %) survival after 2 years and all resprouted within 4 months postfire. In contrast, only 81 % of FS resprouted with only 65 % surviving 2 years postfire. Numerous factors were linked to resprout success: decreased lignotuber exposure and pre-fire vigour (number of pre-fire shoots) were significantly associated with postfire resprouting, while early resprouting (days to first resprout) and growth rate were significant predictors of post-resprout survival. The difference in resprout survival between the FS and OR may also be partially due to phylogenetic differences. These findings confirm the heterogeneity and complexity of postfire resprouting and support the distinction of OS and FS life history types.     

Mortality of resprouting chaparral shrubs after a fire and during a record drought: physiological mechanisms and demographic consequences

We examined postfire regeneration of chaparral shrubs during an intense drought. This study focused on the demography and physiology of shrub species that resprout from a basal lignotuber following fire. We found significant levels of resprout mortality when intense drought occurred in the year following fire during the period of shrub recovery. Three of the seven sampled resprouting species had the greatest or near greatest levels of mortality ever recorded when compared to previous studies. Most shrub mortality occurred during the drought after individuals had resprouted (i.e. individuals survived fire, resprouted and then subsequently died). Physiological measurements of species with high mortality suggested that resprout stems were highly embolized and xylem hydraulic conductivities were close to zero during the peak of the drought. In addition, lignotubers of two of the three species experiencing high mortality were depleted of starch. Population densities of most shrub species declined after the drought compared with their prefire levels, with the exception of one drought tolerant obligate seeding species. Resprouting shrub species may deplete their carbohydrate reserves during the resprouting process, making them particularly vulnerable to drought because of the need to transpire water to acquire the CO₂ that is used to supply energy to a large respiring root system. Drought appears to interact with fire by altering postfire shrub recovery and altering species abundances and composition of chaparral communities.     

大气CO2浓度升高对红桦幼苗根系的影响

应用封闭式生长室系统,研究了CO2浓度升高对红桦(Betula albosinensis)幼苗的根/冠、粗根和细根的干质量、非结构性碳水化合物类含量、碳含量和碳/氮、氮和磷的含量及氮磷吸收量的影响。结果表明:CO2浓度升高使红桦幼苗粗根和细根的干质量增加,同时根/冠值显著升高,表明CO2浓度升高使红桦幼苗生物量向根系的分配增加;与对照相比,粗根的还原糖、蔗糖和总可溶性糖含量显著增加,而在细根中没有显著变化;粗根、细根的淀粉和总的非结构性碳水化合物含量显著增加;CO2浓度升高下粗根和细根的碳含量有升高的趋势但未达到显著水平,同时氮含量降低,碳/氮值升高;氮的吸收量在粗根和细根中均无显著变化。上述结果表明,CO2浓度升高下红桦幼苗根系氮含量下降是由非结构性碳水化合物(主要是淀粉)含量升高和(或)根系生物量增加产生的稀释效应引起的。     

Resource allocation and storage relative to resprouting ability in wind disturbed coastal forest trees

Many plants persist by resprouting after disturbance. However, the benefits of resprouting (survival) may be traded off against height growth and reproduction. Resources (total non-structural carbohydrates—TNC) that could be allocated to growth or reproduction are stored or mobilised to support resprouting. TNC may either be stored by accumulation where availability exceeds the requirements for growth, or by reserve formation when storage is at the expense of growth. Thus, the mechanism of storage and resource allocation may differ between good (R+) and poor (R?) resprouters in response to nutrient availability and disturbance regime. R+ species typically reserve resources to ensure a rapid resprouting response to disturbance. We test whether R+ and R? species in coastal forest, under chronic wind disturbance, differ in growth rates, biomass allocation, leaf traits, water relations and storage of TNC. Seedlings from three confamilial pairs of R+ and R? tree species were subjected to nitrogen addition, water stress and clipping (simulating herbivory) treatments under greenhouse conditions. R? species had greater height growth rates, larger specific leaf area, lower root mass ratio and lower root TNC than R+ species. These differences between R+ and R? species were maintained irrespective of the levels of nitrogen, water and clipping treatments. R+ species did not increase their TNC concentration under nutrient and water stress, indicating that TNC is stored by reserve formation. R+ species appeared to trade-off growth against storage, while R? species did not. In R+ species, reserve formation is likely a bet-hedging strategy against occasional strong selection events in addition to chronic wind stress. By trading off height growth for better resprouting ability, good resprouters may be able to persist at more frequently disturbed sites (e.g., dune crests and windward slopes), while poor resprouters that have faster height growth can dominate less disturbed sites.     

Effect of a strong cold storage induced desiccation on metabolic solutes,stock quality and regrowth,in seedlings of two oak species

Bare-root seedlings of pedunculate oak (Quercus robur L.) and northern red oak (Quercus rubra L.) were lifted in January and stored at 1.8°C, at 82% relative humidity, until their fresh weight declined by 33%. Root growth potential (RGP), fine root electrolyte leakage (REL), fine root water content (RWC), shoot tip water content (SWC), starch and metabolic solute contents in root and shoot, were measured just after lifting and after treatment. Survival of treated seedlings was also assessed in a field trial. RWC, SWC, REL, RGP were dramatically affected by desiccation during cold storage. In both species, root soluble carbohydrate level, inositol level and isocitrate level increased, whereas root starch level and shoot soluble carbohydrate level decreased. In northern red oak, treated seedlings had higher root contents of soluble carbohydrates, inositol and proline than in pedunculate oak. Moreover, treatment induced proline accumulation only in northern red oak roots. These differences could explain why field survival of treated seedlings was significantly better in northern red oak than in pedunculate oak.     

Activities of starch hydrolytic enzymes and starch mobilization in roots of <Emphasis Type="Italic">Caragana korshinskii</Emphasis> following above-ground partial shoot removal

In many resprouting plants, carbohydrates are stored as starch in roots and will be mobilized to support above-ground tissue regrowth after shoot damage. Our objective was to determine how activities of starch hydrolytic enzymes change damage-induced starch mobilization in Caragana korshinskii roots after above-ground tissue loss. Zero percent (control), 30% (30% RSL), 60% (60% RSL) of main shoot length, and 25% (25% RSN), 50% (50% RSN), and 100% (100% RSN) of main shoot number were removed. Compared with control plants, clipping accelerated the reduction of starch in the roots, increased sucrose flux per flower per hour and nectar production per flower per day in 30% RSL, 60% RSL, 25% RSN, and 50% RSN treatments, and improved vegetative growth in 100% RSN treatment. All treatments had similar total nonstructural carbohydrate (TNC) concentrations in leaves, shoots, and stems with the exception of 100% RSN with higher TNC concentration in shoots. Both α-, and β-amylase activities were enhanced by clipping, the former being more strongly correlated with starch degradation in the roots than the latter. The other two possible starch-breaking enzymes, α-glucosidase, and starch phosphorylase showed no significant differences in the activities between treatments. The results suggest that starch degradation in the roots of C. korshinskii was regulated by α-amylase activity and more mobilized starch was used to support vegetative growth in 100% RSN treatment and support sexual reproduction followed by other clipping treatments.     

Resprouting after disturbance: an experimental study with short-lived monocarpic herbs

We experimentally demonstrated the ability of three short-lived monocarpic species to vegetatively regenerate (resprout) from roots after severe disturbance. We assessed the relationship between resprouting ability and (1) timing of injury with respect to life-cycle stage (reproductive vs. vegetative plant), life-history mode (annual vs. winter annual) and phenological stage (flowering vs. fruiting plant), (2) nutrient availability, and (3) disturbance severity (removal of all axillary buds Yes/No). In a chamber experiment with the annual or potentially winter annualRorippa palustris, all injured plants resprouted in all nutrient levels and day-length regimes (day-length regimes simulated conditions of an annual and a winter annual cohort). The number of adventitious buds on roots was positively affected only by injury. The extent of regeneration and amount of regenerated biomass were higher at high nutrient level and long-day regime.     

Seedling vigour and the early growth of transplanted rice (Oryza sativa)

Previous studies suggest that the positive response of transplanted rice (Oryza sativa L.) to nursery fertiliser application was due to increased seedling vigour or possibly to increased nutrient content. This paper presents results of two glasshouse experiments designed to test the hypothesis that seedling vigour was responsible for the response of transplanted seedlings to nursery treatments. The aim of the present study was to explore the concept of seedling vigour of transplanted rice and to determine what plant attributes conferred vigour on the seedlings. Seedling vigour treatments were established by subjecting seedlings to short-term submergence (0, 1 and 2 days/week) in one experiment and to leaf clipping or root pruning and water stress in another to determine their effect on plant growth after transplanting. Submerging seedlings increased plant height but depressed shoot and root dry matter and root:shoot ratio of the seedling at 28 days after sowing. After transplanting these seedlings, prior submergence depressed shoot dry matter at 40 days. Nursery nutrient application increased plant height, increased root and shoot dry matter, but generally decreased root:shoot ratio. Pruning up to 60% of the roots at transplanting decreased shoot and root dry matter, P concentration in leaves at panicle initiation (PI) and straw dry matter and grain yield at maturity. By contrast, pruning 30% of leaves depressed shoot and root dry matter by 30% at PI, and root dry matter and straw and grain yield by 20% at maturity. The combined effects of leaf clipping and root pruning on shoot, root and straw dry matter were largely additive. It is concluded that the response of rice yield to nursery treatments is largely due to increased seedling vigour and can be effected by a range of nutritional as well as non-nutritional treatments of seedlings that increase seedling dry matter, nutrient content, and nutrient concentration. Impairment of leaf growth and to a lesser extent root growth in the nursery depressed seedling vigour after transplanting. However, rather than increasing stress tolerance, seedling vigour was more beneficial when post transplant growth was not limited by nutrient or water stresses.     

Post-fire regeneration strategies of Californian coastal sage shrubs

Summary Regeneration methods for coastal sage srub vegetation after fire were studied in the coastal Santa Monica Mountains of southern California. Six sites were sampled two years after a large fire of fall, 1978. The intensity of fire varied. Foliar cover and flowering incidence were recorded for individuals regenerating by resprouting or from seed. Resprouting plants contributed most to post-fire recovery, comprising 95% of the relative foliar shrub cover; 84% of resprout and 47% of seedling cover had flowered. An ANOVA of reproductive mode and fire intensity indicates that resprout total cover and individual size are significantly greater than those of seedlings, regardless of fire intensity. Among sites the average foliar cover of resprouts exceeded that of seedlings by factors ranging from 9 to 63. All coastal sage species examined resprout, although the potential vigor of resprouting appears to vary widely within genera (e.g. Encelia, Eriogonum, and Salvia) and even within species. In the second growing season following fire seedling density increased due to seeds shed by resprouted shrubs. Most of the cover on these stands of coastal sage scrub is destined to be either crown-sprouted individuals or their progeny.     

Resprouting after disturbance in the short-lived herb Rorippa palustris (Brassicaceae): an experiment with juveniles

The impact of plant age, severity of injury and nutrient levels on the ability to resprout from roots was experimentally assessed in juveniles of the short-lived herb Rorippa palustris (L.) Besser. In a chamber experiment, six cohorts of young plants (1–6 week old) were injured to obtain data on the threshold age for the ability to resprout from roots. We found that plant age was an important factor influencing resprouting ability: injured individuals older than 5 weeks were able to resprout, but not plants younger than 3 weeks. The impact of injury severity (defoliation and removal of axillary buds) and nutrient levels on resprouting ability was assessed on juveniles in a greenhouse experiment. Injury induced growth of new shoots from root buds, while the number of adventitious buds on roots was not influenced by injury. Both injury treatments had a similar effect in this respect, and the amount of regenerated biomass and the extent of regeneration were not different among injury treatments. The number of new shoots produced after injury was higher at the high nutrient level, but the number of formed adventitious buds on roots was not influenced by nutrient level. Nutrient level also influenced the amount of regenerated biomass, but the extent of regeneration (regenerated/removed biomass) was not influenced. The short-lived monocarpic species R. palustris is able to resprout from roots relatively easily. This ability seems to be advantageous in disturbed habitats and this idea is discussed throughout the paper.     

Fire severity and nutrient availability do not constrain resprouting in forest shrubs

Plants often survive disturbances such as fire by resprouting which involves having both protection traits and carbohydrate storage capacity. Protection traits not only act directly to insulate meristems but also prevent combustion of carbohydrate stores. Rapid stem growth also allows replenishment of carbohydrate stores ensuring persistence through another event. Resource availability may, however, constrain the ability to develop resilience to high-severity fires through either nutrient limitation or light limitation. We tested whether fire severity influenced resprouting ability of woody plants in two contrasting environments, low nutrient dry sclerophyll forest and more fertile wet sclerophyll forest. We tested which fire protection and growth traits were associated with resprouting ability (27 species) and resprouting vigour (16 species). Fire severity did not limit the ability of most species to resprout in either forest type. There was no generalized protection syndrome for surviving top kill, but combinations of bud protection and growth together with storage capacity appear to drive resprouting ability. In nutrient-limited forests, low specific leaf area (SLA) may reduce stem growth in resprouters, causing more reliance on bud protection through bark thickness. Conversely, in the more fertile forests, where light becomes limiting with time-since-fire, high SLA appears to increase the capacity for rapid stem growth with less emphasis on developing thicker bark. These different syndromes appear to be adaptive as fire severity did not influence survival in either forest type.     

Is there a cost to resprouting? Seedling growth rate and drought tolerance in sprouting and nonsprouting Ceanothus (Rhamnaceae)

Many woody plant species that depend upon fire-cued seed germination lack the ability to resprout. As the ability to resprout is widely assumed to be the ancestral condition in most plant groups, the failure to sprout is an evolutionary derived trait. Models for the evolutionary loss of sprouting assume a trade-off between seedling success and vegetative resprouting ability of adults. Such models require higher seedling success rates in nonsprouters than in sprouters. On the other hand, there seem to be few a priori reasons why a strong sprouter might not also have highly competitive post-fire seedlings. To test the hypothesis that nonsprouting plants have higher growth rates and/or drought survival, we grew seedlings of Ceanothus tomentosus from sprouting and nonsprouting populations in a common garden experiment. Each of these C. tomentosus populations was paired with a sympatric Ceanothus species that differed in resprouting ability. Sprouters exhibited greater allocation to root carbohydrate storage than did nonsprouters, but overall relative growth rates did not differ. Nonsprouters had earlier onset of flowering. These results provide mixed support for models of a sprouting/nonsprouting allocation trade-off.     

扰动环境中不同刈割方式柠条营养生长补偿的影响

 柠条（Caragana korshinskii）在地上组织破坏后进行补偿性生长,这是重复利用柠条资源的基础,但对 柠条不同刈割方式下营养生长补偿的模式有待探讨。该文通过5种刈割方式：去除主枝长的30%（30%RSL） 、去除主枝长的60%（60%RSL）、去除分枝数的25%（25%RSN）、去除分枝数的50%（50%RSN）和去除分枝 数的100%（100%RSN）来研究柠条的营养生长补偿。结果表明：刈割处理的柠条生物量当年发生了超补偿 ,当年生枝数/枝、当年生枝长、当年生枝生物量/枝、当年生枝生物量/株比对照高。对照、30%RSL和 60%RSL 处理未长出基梢。 100%RSN处理的基梢数/刈割枝、基梢长、单个基梢平均生物量显著高于25%RSN 和50%RSN处理,基梢生物量/株随刈割去除生物量的增加而增加。100%RSN 处理未结果,其它处理果实产量 表现出超补偿或精确补偿。对照处理营养生长和生殖生长均低,其它处理当年生枝生物量与果实产量成显 著负相关。从整个生长季节来看,营养生长主要集中在果实成熟之前。我们认为,100%RSN处理是柠条地上 组织破坏后尽快恢复的合理方式,其当年生生物量远高于其它处理。顶端优势的破坏促使休眠芽的萌发, 根冠比的改变使地上组织获得较多养分和水分,根系储存的碳水化合物的供应是促使刈割柠条营养生长超 补偿的的可能机制,而减少生殖生长对资源的消耗,是100%RSN处理地上生物量尽快恢复的另一重要因素 。     

Fruit production of shrub, Caragana korshinskii, following above-ground partial shoot removal: mechanisms underlying compensation

A number of studies have showed that under some conditions plant may partially, fully or overcompensate for tissue loss, however, the mechanisms underlying compensation are not well understood and still need to be researched. We examined the ability of Caragana korshinskii to compensate for fruit production after above-ground partial shoot removal. Fruit production of 30% main shoot length removal (30% RSL) and 25 and 50% main shoot number removal (25% RSN, 50% RSN) resulted in overcompensation and the response of 60% main shoot length removal (60% RSL) was full compensation. Plants’ responses associated with compensation included (1) greater reproduction efficiency (RA); (2) increased fruit set; (3) decreased fruit abortion; (4) increased seed number per pod; and (5) higher individual seed biomass. These responses may have resulted from more nectar production per flower, more sucrose flux per pod and more sucrose flux per seed of clipped plants, which may in turn have resulted from (1) drawing upon more non-structural carbohydrate (TNC) from roots to supply flower bud development and the flush of new foliage; (2) supplying more photosynthetic assimilation to fruit development owing to increases in leaf-level photosynthetic rates. Increases in leaf-level photosynthetic rates may be caused by more nutrient (nitrogen) and water availability per unit area of resource leaves after clipping.     

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