Interspecific Variation in RGR and the Underlying Traits among 24 Grass Species Grown in Full Daylight

Abstract: A growth analysis was conducted with 24 central European grass species in full daylight to test whether traits underlying interspecific variation in relative growth rate (RGR) are the same in full daylight as they are at lower light, and whether this depends on the ecological characteristics of the studied species, i.e., their requirements with respect to nutrient and light availability.
In contrast to studies with herbaceous species at lower light, net assimilation rate (NAR) contributed more than leaf area ratio (LAR) or specific leaf area (SLA) to interspecific variation in RGR. This was associated with a larger interspecific variation in NAR than found in experiments with lower light. Without the two most shade-tolerant species, however, the contribution of LAR and its components to interspecific variation in RGR was similar or even higher than that of NAR.
Leaf dry matter content correlated negatively with RGR and was the only component of LAR contributing in a similar manner to variation in LAR and RGR. There was a positive correlation between NAR and biomass allocation to roots, which may be a result of nutrient-limited growth. RGR correlated negatively with biomass allocation to leaves. Leaf thickness did not correlate with RGR, as the positive effect of thin leaves was counterbalanced by their lower NAR.
Low inherent RGR was associated with species from nutrient-poor or shady habitats. Different components constrained growth for these two groups of species, those from nutrient-poor habitats having high leaf dry matter content, while those from shady habitats had thin leaves with low NAR.     

Growth Analysis of Sweet Pepper (Capsicum annuum L.): 1. The Influence of Irradiance and Temperature under Glasshouse Conditions in Winter

A growth analysis was carried out with sweet pepper grown ina glasshouse. The plants received natural daylight or additionalillumination applied either during or after the natural photopenod.All irradiance conditions were applied at three temperatureregimes. Additional illumination increased leaf number, leaf area andtotal dry weight. At all temperatures the long-day treatmentsshowed a smaller number of leaves, but a larger leaf area whencompared to the short-day treatments with the same daily radiationsum. A lower temperature progressively reduced leaf area. The derived growth analysis quantities showed strong ontogenetictrends. When comparing both methods of applying additional illuminationhigher mean relative growth rates were observed for the long-daytreatments, especially at the lowest temperature. No differencesin mean net assimilation rate were found, but the short-daytreatments showed a reduced mean leaf area ratio. A lower nighttemperature decreased RGR and NAR but did not affect LAR, alower day temperature increased NAR and decreased LAR. Changesin LAR were largely mediated by changes in specific leaf weight. Capsicum annuum L., sweet pepper, growth analysis, irradiance, temperature     

Regulation of photosynthate partitioning into starch in soybean leaves : response to natural daylight

Studies conducted in controlled environments indicate that daylength affects the proportion of photosynthate stored in leaves as starch or sucrose. To examine the response of partitioning to natural daylight, soybeans (Glycine max [L.] Merr. cv Williams) were grown at 12 different times between May and November in a constant temperature greenhouse without supplemental lighting. Plants were transferred from the greenhouse to a controlled environment chamber at the end of civil twilight at a set developmental stage (expanding seventh trifoliolate leaf, counting acropetally). Net photosynthesis and the accumulation of starch and sugar in fully expanded fourth trifoliolate leaves were determined the following day under standard conditions in the chamber (lights-on synchronized with sunrise). Photosynthesis on a leaf area basis decreased about 10% between midsummer and early autumn. Leaf soluble sugar accumulation was low at all harvests. However, a twofold increase in photosynthate partitioning into starch occurred over the same time period, resulting in an 80% increase in absolute starch accumulation rate. Starch was responsible for about 78% of the increase in leaf dry matter during the light at all harvests, indicating that starch accumulation as affected by prior daylight conditions will alter export of photosynthate during the light period. Photosynthate partitioning into starch was linearly correlated with daylength at harvest, prior average peak solar irradiance, and other parameters that correlated with daylength and solar radiation such as harvested top dry matter. The relation between growth and seasonal changes in daylight (including daylength, irradiance, and light integral) are discussed in relation to photosynthate partitioning under field conditions.     

Growth and dry-mass partitioning in tomato as affected by phosphorus nutrition and light

We studied the effects of phosphorus (P) and light on the physiological and morphological components of growth of young tomato plants (Lycopersicon esculentum Mill. cv. Capita). The importance of dry‐mass partitioning and starch accumulation in explaining the effects of P limitation on growth was examined more closely. Plants were grown at a wide range of exponential P supply rates (between 70 and 320 mg g^?1 d^?1) and one free‐access treatment (1 mm ). Two light levels (70 and 300 µmol m^?2 s^?1) were applied. Growth response coefficients (GRCs) were calculated to address the importance of different growth parameters in explaining relative growth rate (RGR). At both light levels, net assimilation rate (NAR) was more important than leaf area ratio (LAR) in explaining the effects of P on growth as indicated by GRCs. At less severe P limitation, LAR became more important and NAR less important. Dry‐mass partitioning to both roots and leaves played a minor role in determining the effects of P limitation on growth as indicated by low GRCs. The increase in starch at mild P limitation showed that the assimilate supply was not limiting. At severe P limitation, the rate of photosynthesis was decreased, as suggested by the decrease in starch accumulation.     

Growth responses of 15 rain-forest tree species to a light gradient: the relative importance of morphological and physiological traits

1. Growth of seedlings of 15 rain-forest tree species was compared under controlled conditions, at six different light levels (3, 6, 12, 25, 50 and 100% daylight).
2. Most plant variables showed strong ontogenetic changes; they were highly dependent on the biomass of the plant.
3. Growth rate was highest at intermediate light levels (25–50%) above which it declined. Most plant variables showed a curvilinear response to irradiance, with the largest changes at the lowest light levels.
4. There was a consistent ranking in growth between species; species that were fast growing in a low-light environment were also fast growing in a high-light environment.
5. At low light, interspecific variation in relative growth rate was determined mainly by differences in a morphological trait, the leaf area ratio (LAR), whereas at high light it was determined mainly by differences in a physiological trait, the net assimilation rate (NAR).
6. NAR became a stronger determinant of growth than LAR in more than 10–15% daylight. As light availability in the forest is generally much lower than this threshold level, it follows that interspecific variation in growth in a forest environment is mainly owing to variation in morphology.     

光强对两种入侵植物生物量分配、叶片形态和相对生长速率的影响

 比较研究了不同光强下生长的（透光率分别为12.5%、36%、50%、100%）两种入侵性不同的外来种——紫茎泽兰（Eupatorium adenophorum）和兰花菊三七（Gynura sp.）的生物量分配、叶片形态和生长特性。结果表明: 1）两种植物叶片形态对光环境的反应相似。弱光下比叶面积（SLA）、平均单叶面积（MLS）和叶面积比（LAR）较大,随着光强的升高,SLA、MLS、LAR和叶根比（LARMR）降低。2）100%光强下紫茎泽兰叶生物量比（LMR）、叶重分数（LMF）和叶面积指数高于低光强下的值,也高于兰花菊三七,支持结构生物量比（SBR）则相反。强光下紫茎泽兰叶片自遮荫严重,这可能是其表现入侵性的重要原因之一;兰花菊三七分枝较多,避免了叶片自遮荫,较多的分枝利于种子形成对其入侵有利。3）随生长环境光强的升高,两种植物的净同化速率（NAR）、相对生长速率（RGR）和生长对NAR的响应系数均升高（但100%光强下兰花菊三七RGR降低）,平均叶面积比（LARm）和生长对LARm的响应系数均降低,但不同光强下LARm对生长的影响始终大于NAR。4）随着光强的减弱,两种植物都增加高度以截获更多光能,但它们的生物量分配策略不同,紫茎泽兰根生物量比（RMR）降低,SBR增大,而兰花菊三七SBR降低,RMR增大。紫茎泽兰的生物量分配策略更好的反应了弱光环境中的资源变化情况。结论：紫茎泽兰对光环境的适应能力强于兰花菊三七。     

Genotypic difference of potato in carbon budgeting as a mechanism of phosphorus utilization efficiency

Observed genotypic difference in P utilization efficiency in soil grown potatoes led to the present study to investigate possible mechanisms of P utilization efficiency in potato genotypes grown in nutrient solution under three P regimes (low, medium and high). For all genotypes relative growth rate (RGR), leaf P content, net assimilation rate (NAR) and leaf area ratio (LAR) increased while P utilization efficiency and leaf starch content decreased at the two higher P regimes compared to the low P regime. The P-efficient genotypes CGN 17903 and CIP 384321.3 had higher RGR compared to the P-inefficient genotypes CGN 22367 and CGN 18233, which resulted from enhanced NAR rather than from LAR. Net photosynthetic rate was similar for all genotypes. However, for P-inefficient genotype CGN 22367, the lower NAR could be explained by increased leaf dark respiration. For P-inefficient genotype CGN 18233 we speculate that increased carbon cost of root respiration or exudation or both, caused low NAR, since leaf dark respiration of this genotype was similar to that of P-efficient genotypes.     

Effects of irradiance on relative growth rates, net assimilation rates, and leaf area partitioning in cotton and three associated weeds

Cotton (Gossypium hirsutum L. var. `Stoneville 213'), velvetleaf (Abutilon theophrasti Medic.), redroot pigweed (Amaranthus retroflexus L.), and hemp sesbania (Sesbania exaltata [Raf.] Cory) were grown in a controlled environment room at 31/25 C day/night temperature and three irradiances: 90, 320, and 750 μeinsteins meter⁻² second⁻¹. From total dry weights and leaf areas determined at intervals during the first exponential phase of growth, we used mathematical growth analysis techniques to calculate net assimilation rates (NAR), relative growth rates (R_w), relative leaf area expansion rates (R_a), leaf area partition coefficients (LAP), and leaf area ratios (LAR). In all four species, R_w, R_a, and NAR decreased with decreasing growth irradiance, while LAP and LAR increased. Within each species, R_w was positively correlated with NAR but negatively correlated with LAP and LAR. In comparisons among the four species within each growth irradiance, R_w was positively correlated with LAP. We discuss the relationship between LAP and LAR and show that LAP = (R_a/R_w) (LAR).     

Free proline accumulation in drought-stressed plants under laboratory conditions

Summary Free proline accumulation was measured in leaves of intact sorghum (Sorghum bicolor L. cv. Pioneer 846) and soybean (Glycine max. L. cv. Calland) grown in growth chambers and subjected to ‘normal’ drought stress. Stomatal diffusive resistance and leaf water potential were used to determine the degree of stress at the time of proline analysis. Free proline did not accumulate markedly in either species until each was severely stressed, indicating that proline is not a sensitive indicator of drought stress. Free proline accumulated under less stress in soybean than in sorghum. Since soybean is less drought resistant than sorghum, proline accumulation may be an indicator of drought resistance or susceptibility. re]19731003     

New seed morphological features in Moehringia L. (Caryophyllaceae) and their taxonomic and ecological significance

Abstract

Relative growth rate (RGR) is a fundamental trait for comparative plant ecology but cannot be measured in situ, leading to problems in interpreting vegetation function. However, the components of RGR (net assimilation rate (NAR), leaf area ratio (LAR), leaf weight ratio (LWR), and specific leaf area (SLA)) can be calculated for wild plants from morphological measurements (leaf area, leaf dry mass, whole plant dry mass), which potentially reflect RGR. Seeds of 19 species from Italian prealpine calcareous grasslands were collected and seedlings were cultivated under controlled conditions. RGR, NAR, LAR, LWR and SLA were analysed. The results demonstrated that RGR was positively correlated with SLA and LAR (p < 0.01). Furthermore, LAR was positively correlated with LWR and negatively with NAR (p < 0.05). Monocotyledons showed significantly higher LAR, LWR and NAR than dicotyledons, as the latter allocated a greater proportion of biomass to stems, but RGR and SLA showed no such phylogenetic constraint. Therefore SLA is the most reliable indicator of RGR in ecological and functional surveys of prealpine calcareous grasslands, and has the additional advantage that it can be measured from leaf material alone. Lower mean RGR and SLA for calcareous grassland species suggests that this vegetation is less likely to recover from the effects of disturbance than meadows and dry meadows.     

Osservazioni Preliminari Sull'ecologia di Semenzali di Rovere (Quercus Petraea (Matt.) Liebl.)

Abstract

Preliminary observations on the ecology of sessile oak (Quercus petraea (Matt.) Liebl.) seedlings.—The responses of sessile oak seedlings to four different light intensities and to increasing drought (three soil moisture contents) have been investigated in a nursery experiment in order to improve the knowledge of the ecology of this species in the Mediterranean area. Data have been compared with observations on natural regeneration established in different shade and soil moisture conditions in a sessile oak stand. The results of the two experiments showed that low light intensities (transmittance <5%) did not limit the seedlings establishment but induced lowest growth rates, while in full daylight seedlings mortality was very high (61%). In accordance with the observations in natural conditions, seedlings showed the best performances at intermediate light levels (50–80% of full daylight) and with increasing soil moisture: mortality was very low and average height increment, dry weight, relative growth rate (RGR) and net assimilation rate (NAR) showed the highest values. The differences in specific leaf area (SLA), leaf area ratio (LAR), root/stem ratio enabled to discuss the sessile oak seedlings ecology and adaptations to different light intensities and soil moisture content.     

Optimal leaf area indices in C3 and C4 mono- and dicotyledonous species at low and high nitrogen availability

From an analytical model it was shown that for a given total amount of nitrogen in the canopy, there exists an optimal leaf area index (LAI), and therefore an optimal average leaf introgen content, at which canopy photosynthesis is maximal. If the LAI is increased above this optimum, increased light interception will not compensate for reduction in photosynthetic capacity of the canopy resulting from reduced leaf nitrogen contents. It was further derived from the model that the value of the optimal LAI increases with the photosynthetic nitrogen use efficiency (PNUE) and decreases with the canopy extinction coefficient for light (K_L) and incident photon flux density (PFD) at the top of the canopy. These hypotheses were tested on dense stands of species with different photosynthetic modes and different architectures. A garden experiment was carried out with the C₄ monocot sorghum ( Sorghum bicolor [L.] Moensch cv. Pioneer), the C₃ monocot rice ( Oryza sativa L. cv. Araure 4), the C₄ dicot amaranth ( Amaranthus cruentus L. cv. K113) and the C₃ dicot soybean ( Glycine max [L.] Merr. cv. Williams) at two levels of nitrogen availability.
The C₄ species had higher PNUEs than the C₃ species while the dicots formed stands with higher extinction coefficients for light and had lower PNUEs than the monocots. The C₄ and monocot species were found to have formed more leaf area per unit leaf nitrogen (i.e., had lower leaf nitrogen contents) than the C₃ and dicot species, respectively. These results indicate that the PNUE and the extinction coefficient for light are important factors determining the amount of leaf area produced per unit nitrogen as was predicted by the model.     

Assimilation Rate and Growth of Lolium Populations in the Glasshouse in Contrasting Light Intensities

Using growth-analysis techniques, the variation in relativegrowth-rate (RGR) and its components, net assimilation rate(NAR), and leaf-area ratio (LAR), was examined in 18 populationsof L. perenne, six of L. multiflorum, and two hybrid cultivarsfrom contrasting climatic and agronomic origins, grown at lowand high light intensities in the glasshouse. Significant differences between populations were found for RGR,NAR, and LAR at both light intensities. At both intensitiesthe annual or biennial multiflorum group had a greater LAR anda lower specific leaf weight and chlorophyll content than theperennial perenne group. At the low intensity this was compensatedby a greater NAR in the perenne group, with no resultant differencein RGR. At the high intensity there was no difference betweenthe groups in NAR, and hence a greater RGR in the multiflorumgroup. Within the perenne and multiflorum groups, at both light intensities,the variation between populations in RGR was based on differencesin NAR rather than in LAR. There was no regular correlationof NAR with either specific leaf weight, or chlorophyll contentat either light intensity, though at low light intensity itwas significantly correlated with shoot-root ratio.     

Photosynthate Partitioning into Starch in Soybean Leaves: I. Effects of Photoperiod versus Photosynthetic Period Duration

Photosynthesis, photosynthate partitioning into foliar starch, and translocation were investigated in soybean plants (Glycine max (L.) Merr. cv. Amsoy 71), grown under different photoperiods and photosynthetic periods to determine the controls of leaf starch accumulation. Starch accumulation rates in soybean leaves were inversely related to the length of the daily photosynthetic period under which the plants were grown. Photosynthetic period and not photoperiod per se appears to be the important factor. Plants grown in a 14-hour photosynthetic period partitioned approximately 60% of the daily foliar accumulation into starch whereas 7-hour plants partitioned about 90% of their daily foliar accumulation into starch. The difference in starch accumulation resulted from a change in photosynthate partitioning between starch and leaf residual dry weight. Residual dry weight is defined as leaf dry weight minus the weight of total nonstructural carbohydrates. Differences in photosynthate partitioning into starch were also associated with changes in photosynthetic and translocation rates, as well as with leaf and whole plant morphology. It is concluded that leaf starch accumulation is a programmed process and not simply the result of a limitation in translocation.     

Net assimilation rate and relative nitrogen assimilation rate in relation to the dry matter production of alfalfa cultivars

Summary Twelve alfalfa cultivars inoculated with an indigenous strain (RM9) ofRhizobium meliloti, were compared for their seedling morphological characters, and growth characters, including net assimilation rate (NAR), relative growth rate (RGR), leaf area ratio (LAR) and relative nitrogen assimilation rate (R_N). Highly significant differences were obtained between cultivars for most characters.Simple correlation showed that NAR influenced RGR (r=0.91) more than leaf area ratio (LAR) (r=–0.44), and that most characters measured were highly correlated with seedling dry weight. Factor analysis showed that NAR, RGR and R_N contributed 25% of the total variation in the dependence structure. The grouping indicated that the higher the NAR and R_N the greater was the RGR. Path-coefficient analysis showed that NAR had more important direct and indirect effects than R_N in dry matter accumulation. The relationship implied that selection for plants with high NAR, or high efficiency in converting light energy to dry matter production could contribute greater N₂ fixation in alfalfa.     

Variation in relative growth rate of 20 Aegilops species (Poaceae) in the field: The importance of net assimilation rate or specific leaf area depends on the time scale

Field experiments reporting the relative growth rate (RGR) patterns in plants are scarce. In this study, 22 herbaceous species (20 Aegilops species, Amblyopyrum muticum and Triticum aestivum) were grown under field conditions to assess their RGR, and to find out if the differences in RGR amongst species were explained by morphological or physiological traits. Plants were cultivated during two months, and five harvests (every 13–19 days) were carried out. Factors explaining between-species differences in RGR varied, depending on whether short (13–19 days) or longer periods (62 days) were considered. RGR for short periods (4 growth periods of 13–19 days each) showed a positive correlation with net assimilation rate (NAR), but there was no significant correlation with leaf area ratio (LAR) (with the exception of the first growth period). In contrast, when growth was investigated over two months, RGR was positively correlated with morphological traits (LAR, and specific leaf area, SLA), but not with physiological traits (NAR). A possible explanation for these contrasting results is that during short growth periods, NAR exhibited strong variations possibly caused by the variable field conditions, and, consequently NAR mainly determined RGR. In contrast, during a longer growth period (62 days) the importance of NAR was not apparent (there was no significant correlation between RGR and NAR), while allocation traits, such as LAR and SLA, became most relevant.     

Waterlogging injury in sorghum seedlings and their kinetin effected rapid recovery

Short term waterlogging affected the growth of sorghum seedlings as indicated by a high mortality rate of the seedlings and a decrease in the shoot and root biomass, net assimilation rate (NAR), leaf area ratio (LAR), relative growth rate (RGR), total photosynthetic area and rate of photosynthesis. Plant resistance to the stress was exhibited by decreased stomatal index (SI), rate of transpiration and rate of respiration. When the stressed plants were exposed to normal conditions, the recovery of the seedlings was rapid and was enhanced by foliar application of kinetin at 5 mgl^-1.     

The response of growth rate to temperature in seedlings of several Allium crop species

Relative growth rates (R_w), leaf area ratios (LAR), net assimilation rates (NAR), and leaf weight ratios (LWR) were estimated for seedlings of onion, leek, Japanese bunching onion and chives grown at constant temperatures of 10, 12, 15, 19, 23, 27 and 31 C. R_w was greatest at 19 C in Japanese bunching onion and chives, and at 27 C in onions and leeks. R_w increased approximately linearly between 10 and 19 C in all species; between 19 and 27 C it either increased further, decreased or remained constant depending on species but at 31 C R_w was lower than the maximum in all cases. LAR's were greatest at 27 C in onions and leeks and at 23 C in the other two species. Between 10 and 19 C both LAR and NAR increased in all species. Between 19 and 27 C increases in LAR were compensated for by decreases in NAR with the result that R_w showed little change with temperature. Averaging over all temperatures the species were ranked in order of decreasing R_w, NAR and LWR as onion, leek, Japanese bunching onion and chives. For LAR this ordering was reversed with respect to onions and leeks. Plant to plant variability increased in the order onion, leek, Japanese bunching onion then chives. Although onions were the fastest growing of the Alliums tested their R_w was lower than that of several other vegetable species for which R_w was similarly measured.     

Phenotypic plasticity of lianas in response to altered light environment

The growth, morphology and biomass allocation of 11 liana species (six light-demanding and five shade-tolerant) were investigated by growing plants in three contrasting light environments (i.e., field, forest edge and forest interior). Our objectives were to determine: (1) changes in plant traits at the species level; and (2) differences in light-demanding and shade-tolerant species in response to altered light environment. We found that all seedlings of liana species increased in total biomass, total leaf area, relative growth rate (RGR), net assimilation rate (NAR), height, basal diameter, root length, leaf number, root mass/total plant mass (RMR) and root-to-shoot dry biomass (R/S ratio), and decreased in leaf area ratio (LAR), specific leaf area (SLA), leaf size, stem mass-to-total plant mass ratio (SMR) and leaf mass-to-total plant mass ratio (LMR) with increasing light availability. Under the three light environments, the two types of species differed significantly in total biomass, total leaf area, RGR, NAR, LAR, SLA and leaf number, and not in leaf area. Only light-demanding species differed significantly in height, root length, basal diameter, RMR, SMR, LMR and R/S ratio. The mean plasticity index of growth and biomass allocation were relatively higher than the morphological variables, with significant differences between the two groups. Our results showed that liana species respond differently to changing light environments and that light-demanding species exhibit higher plasticity. Such differences may affect the relative success of liana species in forest dynamics.     

Chloroplast Structure and Starch Grain Accumulation in Leaves That Received Different Red and Far-Red Levels during Development

An important step in understanding influence of growth environment on carbon metabolism in plants is to gain a better understanding of effects of light quality on the photosynthetic system. Electron microscopy was used to study chloroplast ultrastructure in developing and fully expanded leaves of tobacco (Nicotiana tabacum L. cv Burley 21). Brief exposures to red or far-red light at the end of each day during growth under controlled environments influenced granum size, granum number and starch grain accumulation in chloroplasts, and the concentration of sugars in leaf lamina. Far-red-treated leaves had chloroplasts with more but smaller grana than did red-treated leaves. Red light at the end of the photosynthetic period resulted in more and larger starch grains in the chloroplasts and a lower concentration of sugars in leaves. Chloroplast ultrastructure and starch grain accumulation patterns that were initiated in the expanding leaves were also evident in the fully expanded leaves that received the treatment during development. It appears that the phytochrome system in the developing leaves sensed the light environment and initiated events which influenced chloroplast development and partitioning of photosynthate to adapt the plant for better survival under those environmental conditions.