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.     

Interspecific correlates of plasticity in relative growth rate following a decrease in nitrogen availability

Background and Aims

Nitrogen availability varies greatly over short time scales. This requires that a well-adapted plant modify its phenotype by an appropriate amount and at a certain speed in order to maximize growth and fitness. To determine how plastic ontogenetic changes in each trait interact and whether or not these changes are likely to maximize growth, ontogenetic changes in relative growth rate (RGR), net assimilation rate (NAR), specific leaf area (SLA) and root weight ratio (RWR), before and after a decrease in nitrogen supply, were studied in 14 herbaceous species.

Methods

Forty-four plants of each species were grown in hydroponic culture under controlled conditions in a control treatment where the supply of nitrogen remained constant at 1 mm, and in a stress treatment where the nitrogen supply was abruptly decreased from 1 to 0·01 mm during the growth period.

Key Results and Conclusions

In the treatment series, and in comparison with the control, NAR and RGR decreased, RWR increased, and SLA did not change except for the timing of ontogenetic change. Species having greater increases in the maximum rate of change in RWR also had smaller reductions in RGR; plasticity in RWR is therefore adaptive. In contrast, species which showed a greater decrease in NAR showed stronger reductions in RGR; plasticity in NAR is therefore not adaptive. Plasticity in RGR was not related to plasticity in SLA. There were no significant relationships among the plasticities in NAR, RWR or SLA. Potentially fast-growing species experienced larger reductions in RGR following the nitrogen reduction. These results suggest that competitive responses to interspecific competition for nitrogen might be positively correlated with the plasticity in the maximum rate of change in RWR in response to a reduction in nitrogen supply.     

Leaf area ratio and net assimilation rate of 24 wild species differing in relative growth rate

Summary Which factors cause fast-growing plant species to achieve a higher relative growth rate than slow-growing ones? To answer this question 24 wild species were grown from seed in a growth chamber under conditions of optimal nutrient supply and a growth analysis was carried out. Mean relative growth rate, corrected for possible ontogenetic drift, ranged from 113 to 356 mg g^–1 day^–1. Net assimilation rate, the increase in plant dry weight per unit leaf area and unit time, varied two-fold between species but no correlation with relative growth rate was found. The correlation between leaf area ratio, the ratio between total leaf area and total plant weight, and relative growth rate was very high. This positive correlation was mainly due to the specific leaf area, the ratio between leaf area and leaf weight, and to a lesser extent caused by the leaf weight ratio, the fraction of plant biomass allocated to the leaves. Differences in relative growth rate under conditions of optimum nutrient supply were correlated with the soil fertility in the natural habitat of these species. It is postulated that natural selection in a nutrient-rich environment has favoured species with a high specific leaf area and a high leaf weight ratio, and consequently a high leaf area ratio, whereas selection in nutrient-poor habitats has led to species with an inherently low specific leaf area and a higher fraction of root mass, and thus a low leaf area ratio.     

Modelling the relative growth rate as a function of plant nitrogen concentration

The relationship between the relative growth rate (RGR) and the nitrogen concentration of the whole plant (PNC) was analyzed by using experimentally determined relations (1) between the PNC and the fraction of dry matter (LWR) and nitrogen in leaves, (2) between the specific leaf area (SLA) and the leaf nitrogen concentration (LNC) and (3) between the net assimilation rate (NAR) and the LNC on an area basis. A strong dependence of RGR on nitrogen concentration resulted from the increase in NAR, LWR and SLA with increasing PNC. A curvilinear relationship between RGR and PNC gave an optimum curve for nitrogen productivity against PNC.     

Differences in developmental plasticity and growth rate among drought-resistant and susceptible cultivars of durum wheat (Triticum turgidum L. var. durum)

Understanding how growth and development of durum wheat cultivars respond to drought could provide a basis to develop crop improvement programmes in drought-affected tropical and subtropical countries. A greenhouse experiment was conducted to study the responses of five durum wheat cultivars to moisture stress at different developmental phases. Phenology, total dry matter (TDM), relative growth rate (RGR), leaf area ratio (LAR), net assimilation rate (NAR), leaf weight ratio (LWR), specific leaf area (SLA) and shoot:root ratio were compared. Pre-anthesis moisture stress delayed phenological development, whereas post-anthesis moisture stress accelerated it. TDM accumulation rate was different between drought-resistant and susceptible cultivars. RGR and its components changed with age and moisture availability. Drought-resistant cultivars had a high RGR in favourable periods of the growing season and a low RGR during moisture stress. In contrast, the drought-susceptible cultivar (Po) showed an opposite trend. LAR explained the differences in RGR (r=0.788) best, whereas the relationship between NAR and RGR was not significant. Even though both LWR and SLA were important factors determining the potential growth rate, LWR was of major importance to describe cultivar differences in LAR, and consequently in RGR. The drought-resistant cultivars Omrabi-5 and Boohai showed vigorous root development and/or a low shoot:root ratio. It is concluded that biomass allocation is the major factor explaining variation in RGR among the investigated durum wheat cultivars.     

Relative growth rate and biomass allocation in ten woody species with different leaf longevity using phylogenetic independent contrasts (PICs)

In this study, we compare the relative growth rate (RGR) and biomass allocation of 10 woody species (5 deciduous and 5 evergreen) from the Mediterranean region using phylogenetic independent contrasts (PICs) to test if these two functional groups differ in these traits. In general, the results were similar when using PICs or without taking into account phylogenetic relations. Deciduous species had a higher RGR than evergreen species, due to the higher net assimilation rate (NAR). Deciduous species had a higher specific leaf area (SLA) but a lower leaf mass ratio (LMR), resulting in a similar LAR for deciduous and evergreen species (LAR = SLA x LMR). In some cases, the use of PICs revealed patterns that would not have appeared if phylogeny had been overlooked. For example, there was no significant correlation between RGR and final dry mass (after 4 months of growth) but PICs revealed that there was a positive relation between these two variables in all deciduous-evergreen pairs. In general, RGR decreased with time and this temporal variation was due primarily to NAR variations (r = 0.79, p < 0.01), and also to variations in LAR (r = 0.69, p < 0.05). Considering the phylogeny, the only variable constantly different for all deciduous-evergreen pairs was SLA. This result, and the fact that SLA was the best correlated variable with RGR (r = 0.81, p < 0.01), reinforce the value of SLA as a variable closely associated to growth and to the functional groups (deciduous vs. evergreen).     

Responses of photosynthesis,leaf conductance and growth to different salinities in three coastal dune plants

The effects of soil-water salinity on growth and photosynthesis of three coastal dune plants were examined by salt-treatment in order to clarify the causal relationship between salinity and plant distribution in a dune habitat. Plants were cultivated hydroponically at three salinity levels: 0, 10 and 100 mM NaCl. With the 100 mM salt treatment,Calystegia soldanella (C₃ species) had the highest relative growth rate (RGR) (0.085 g g⁻¹ d⁻¹), followed byCarex kobomugi (C₃) (0.066), andIschaemum anthephoroides (C₄) (0.060). This order coincides with the distribution pattern of the three species on coastal dunes;Calystegia soldanella is generally distributed in more seaward areas whereasI. anthephoroides occurs further inland. The order of RGR was determined exclusively by leaf area ratio (LAR) among the three species. Due to its C₄ pathway,I. anthephoroides had higher net photosynthetic rate (Pn) and net assimilation rate (NAR) than the two C₃ plants at all NaCl concentrations, despite its low RGR. This apparent discrepancy is explainable by differences of LAR among the three species; LAR ofI. anthephoroides was lowest, and about half that ofCalystegia soldanella. These results suggest that LAR is one of the main determinants of salt tolerance based on RGR, whereas Pn or NAR may not be significant. This article is dedicated to Professor Hideo Iwaki, University of Tsukuba, in appreciation of the sincere encouragement he has given to the authors.     

Plasticity in relative growth rate and its components following a change in irradiance

A total of 244 plants from two species, Lythrum salicaria and Epilobium glandulosum, were grown individually in hydroponic sand culture from seed for 36 d. Until day 27 all plants experienced an irradiance of 550 μmol m^?2 s^?1 PFD and on day 27 half of the plants were subjected to a neutral shade treatment in which irradiance was reduced to 100 μmol m^?2 s^?1 photon fluy density (PFD). Measures of relative growth rate, net assimilation rate, specific leaf area, biomass partitioning to leaves, roots, structural tissues (i.e. stems, petioles and inflorescences) and tissue density were obtained from intensive harvests three or four times per day. The shade treatment caused an immediate decrease in relative growth rate and net assimilation rate. Within hours the specific leaf area of the shaded plants increased and leaf tissue density decreased, thus partially offsetting the decrease in relative growth rate. Biomass partitioning was not affected.     

Consequences of phenotypic plasticity vs. interspecific differences in leaf and root traits for acquisition of aboveground and belowground resources

Trade-offs between acquisition capacities for aboveground and belowground resources were investigated by studying the phenotypic plasticity of leaf and root traits in response to different irradiance levels at low nutrient supply. Two congeneric grasses with contrasting light requirements, Dactylis glomerata and D. polygama, were used. The aim was to analyze phenotypic covariation in components of leaf area and root length in response to above- and belowground resource limitation and the consequences of this variation for resource acquisition and plant growth. At intermediate shading (30 and 20% of full sunlight) the plants were able to maintain their total root length, despite a strongly increased total leaf area and a reduced biomass allocation to roots. This was associated with an unaltered or slightly increased nutrient uptake and growth. At 5.5% relative irradiance, growth was severely reduced, especially in the shade-tolerant D. polygama. The results show that constraints on acquisition capacities for aboveground and belowground resources, caused by biomass allocation, may be alleviated by plasticity in other traits such as tissue-mass density and thickness of roots and leaves. The results also suggest different adaptive constraints for phenotypic plasticity and for genetically determined interspecific variation. Phenotypic plasticity tends to maximize resource acquisition and growth rate in the short term, whereas the higher tissue-mass density and the longer leaf life-span of shade-tolerant species indicate reduced loss rates as a more advantageous species-specific adaptation to shade in the long term.     

Supplemental manganese improves the relative growth, net assimilation and photosynthetic rates of salt-stressed barley

Previous results in our laboratory indicated that a reduced Mn concentration in the leaves of barley was highly correlated with the reduced relative growth and net assimilation rates of salt-stressed plants. If Mn deficiency limits the growth of salt-stressed barley, then increasing leaf Mn concentrations should increase growth. In the present study, the effect of supplemental Mn on the growth of salt-stressed barley ( Hordeum vulgare L. cv. CM 72) was tested to determine if a salinity-induced Mn deficiency was limiting growth. Plants were salinized with 125 mol m⁻³ NaCl and 9.6 mol m⁻³ CaCl₂. Supplemental Mn was applied in 2 ways: 1) by increasing the Mn concentration in the solution culture and 2) by spraying Mn solutions directly onto the leaves. Growth was markedly inhibited at this salinity level. Dry matter production was increased 100% in salt-stressed plants treated with supplemental Mn to about 32% of the level of nonsalinized controls. The optimum solution culture concentration was 2.0 mmol m⁻³, and the optimum concentration applied to the leaves was 5.0 mol m⁻³. Supplemental Mn did not affect the growth of control plants. Further experiments showed that supplemental Mn increased Mn concentrations and uptake to the shoot. Supplemental Mn increased the relative growth rate of salt-stressed plants and this increase was attributed to an increase in the net assimilation rate; there were no significant effects on the leaf area ratio. Supplemental Mn also increased the net photosynthetic rate of salt-stressed plants. The data support the hypothesis that salinity induced a Mn deficiency in the shoot, which partially reduced photosynthetic rates and growth.     

Growth temperature influences the underlying components of relative growth rate: an investigation using inherently fast- and slow-growing plant species

We examined the effect of growth temperature on the underlying components of growth in a range of inherently fast‐ and slow‐growing plant species. Plants were grown hydroponically at constant 18, 23 and 28 °C. Growth analysis was conducted on 16 contrasting plant species, with whole plant gas exchange being performed on six of the 16 species. Inter‐specific variations in specific leaf area (SLA) were important in determining variations in relative growth rate (RGR) amongst the species at 23 and 28 °C but were not related to variations in RGR at 18 °C. When grown at 18 °C, net assimilation rate (NAR) became more important than SLA for explaining variations in RGR. Variations in whole shoot photosynthesis and carbon concentration could not explain the importance of NAR in determining RGR at the lower temperatures. Rather, variations in the degree to which whole plant respiration per unit leaf area acclimated to the different growth temperatures were responsible. Plants grown at 28 °C used a greater proportion of their daily fixed carbon in respiration than did the 18 and 23 °C‐grown plants. It is concluded that the relative importance of the underlying components of growth are influenced by growth temperature, and the degree of acclimation of respiration is of central importance to the greater role played by NAR in determining variations in RGR at declining growth temperatures.     

Growth and photosynthetic response of three soybean cultivars to simultaneous increases in growth temperature and CO2

Three soybean ( Glycine max L. Merr.) cultivars (Maple Glen, Clark and CNS) were exposed to three CO₂ concentrations (370, 555 and 740 μmol mol⁻¹) and three growth temperatures (20/15°, 25/20° and 31/26°C, day/night) to determine intraspecific differences in single leaf/whole plant photosynthesis, growth and partitioning, phenology and final biomass. Based on known carboxylation kinetics, a synergistic effect between temperature and CO₂ on growth and photosynthesis was predicted since elevated CO₂ increases photosynthesis by reducing photorespiration and photorespiration increases with temperature. Increasing CO₂ concentrations resulted in a stimulation of single leaf photosynthesis for 40–60 days after emergence (DAE) at 20/15°C in all cultivars and for Maple Glen and CNS at all temperatures. For Clark, however, the onset of flowering at warmer temperatures coincided with the loss of stimulation in single leaf photosynthesis at elevated CO₂ concentrations. Despite the season-long stimulation of single leaf photosynthesis, elevated CO₂ concentrations did not increase whole plant photosynthesis except at the highest growth temperature in Maple Glen and CNS, and there was no synergistic effect on final biomass. Instead, the stimulatory effect of CO₂ on growth was delayed by higher temperatures. Data from this experiment suggest that: (1) intraspecific variation could be used to select for optimum soybean cultivars with future climate change; and (2) the relationship between temperature and CO₂ concentration may be expressed differently at the leaf and whole plant levels and may not solely reflect known changes in carboxylation kinetics.     

Relating root structure and anatomy to whole-plant functioning in 14 herbaceous Mediterranean species

This study investigated the relationships between root structure and anatomy and whole-plant functioning in herbaceous species. Fourteen annual and perennial species representative of a Mediterranean old-field succession were grown in monocultures in a common-garden experiment. Whole-plant functioning was assessed by inherent relative growth rate (RGR(max)), measured in standardized conditions, and maximum height (H(max)). Root tissue density (TMD(r)), considered as a major component of root structure, was measured on roots harvested within in-growth cores. Anatomical characteristics were analysed on cross-sectional areas (CSA). TMD(r) was correlated positively with H(max) and negatively with RGR(max). Root CSA explained interspecific variation in H(max) but not that in TMD(r) and RGR(max). Root xylem CSA and xylem proportion in root CSA were positively correlated with TMD(r) and H(max) and negatively with RGR(max). Mean xylem vessel CSA did not account for variations in TMD(r), H(max) and RGR(max). These results suggested that RGR(max) and H(max) are constrained by opposite root structural and anatomical traits, which have potential links with hydraulic conductance, support and longevity.     

Intra-specific variation in relative growth rate: impact on competitive ability and performance of Lychnis flos-cuculi in habitats differing in soil fertility

Plant species from unproductive or adverse habitats are often characterized by a low potential relative growth rate (RGR). Although it is generally assumed that this is the result of selection for specific trait combinations that are associated with a low rate of net biomass accumulation, few studies have directly investigated the selective (dis-)advantage of specific growth parameters under a set of different environmental conditions. Aim of the present study was to quantify the impact of inherent differences in growth parameters among phenotypes of a single plant species, Lychnis flos-cuculi, on their performance under different soil nutrient conditions. Growth analysis revealed significant variation in RGR among progeny families from a diallel cross between eight genotypes originating from a single population. Differences in RGR were due to variation in both leaf area ratio (LAR) and in net assimilation rate (NAR). A genetic trade-off was observed between these two components of growth, i.e. progeny families with high investment in leaf area had a lower rate of net biomass accumulation per unit leaf area. The degree of plasticity in RGR to nutrient conditions did not differ among progeny families. Inherent differences in growth parameters among progeny families had a significant impact on their yield in competition with Anthoxanthum odoratum and Taraxacum hollandicum. In nutrient-rich conditions, progeny families with an inherently high leaf weight ratio (LWR) achieved higher yield in competition, but variation in this trait could not explain differences in competitive yield under nutrient-poor conditions. Inherent differences in growth parameters among progeny families were poorly correlated with differences in survival and average rosette biomass (a good predictor of fecundity) among these progeny families sown in four field sites along a natural gradient of soil fertility. In the more productive sites none of the growth parameters was significantly correlated with rosette biomass, but in the least productive site progeny families with an inherently high specific leaf area (SLA) tended to produce smaller rosettes than low-SLA families. These results are consistent with the view that a selective advantage may accrue from either high or low values of individual RGR components, depending on habitat conditions, and that the selective advantage of low trait values in nutrient-poor environments may results in indirect selection for low RGR in these habitats.     

Relative addition rate and external concentration; Driving variables used in plant nutrition research

Abstract An increasing literature accounting for various types of experiments indicates that far lower external nutrient concentrations are required by plants than is usually thought to be the case. It is concluded that the ion uptake capacity of the roots, as described by the carrier concept, is high compared to that required for maintenance of the internal concentration. Serious errors in experimental conclusions are associated with insufficient and constant nutrient addition rates. The main errors are caused by non-steady states of the plants both with regard to the internal nutrient concentrations and the relative growth rate. A dynamic concept has been proposed for direct use as the treatment variable within the range of sub-optimum nutrition. The nutritional factor is expressed as a flow, the relative nutrient addition rate in laboratory studies and the nutrient flux density in the field. The experimental use of the relative addition rate has led to steady-state nutrient status and relative growth rate and the interpretation of plant responses which differ fundamentally from accepted views. Thus, for instance, deficiency symptoms disappear, as in natural conditions, when the internal nitrogen concentration is stable, independent of level. The nutrition/growth relationships are very different from those observed when external concentration is varied. The regression line of relative growth rate on relative addition rate passes near to the origin at an angle close to 45 to the axes, which implies that the obtained relative growth rate approximates closely the treatment variable. A striking example of observed differences is the positive effect on nitrogen fixation exerted by high relative nitrogen addition rates compared to the well-known negative effect of increasing external nitrogen concentration. The application of fertilizer on the basis of the nutrient flux density concept provides the possibility of supplying fertilizers corresponding to the consumption potential of the vegetation and to the natural flux density resulting from mineralization in the soil. Nitrogen utilization is high under such conditions and the resulting feedback of nutrition on the mineralization rate suggests that there will be a long-term increase in fertility.     

Growth analysis of UV-B-irradiated cucumber seedlings as influenced by photosynthetic photon flux source and cultivar

A growth analysis was made of ultraviolet-B (UV-B)-sensitive (Poinsett) and insensitive (Ashley) cultivars of Cucuumis satives L. grown in growth chambers at 600 μmol m⁻² s⁻¹ of photosynthetic photon flux (PPF) provided by red- and far-red-deficient metal halide (MH) or blue- and UV-A-deficient high pressure sodium/deluxe f HPS/DX) lamps. Plants were irradiated 6 h daiiy with 0.2 f-UV-B) or 18.2 C+UV-B) kJ m⁻² day⁻¹ of biologically effective UV-B for 8 or 15 days from time of seeding. In general, plants given supplemental UV-B for 15 days showed lower leaf area ratio (LARs, and higher specific leaf mass (SLM) mean relative growth rate (MRGR) and net assimilation rate (NAR) than that of control plants, but they showed no difference in leaf mass ratio (LMR), Plants grown under HPS/DX lamps vs MH lamps showed higher SLM and NAR. lower LAR and LMR. hut no difference in MRGR. LMR was the only growth parameter affected by cultivar: at 15 days, it was slightly greater in Poinsett than in Ashley. There were no interactive effects of UV-B. PPF source or cultivar on any of the growth parameters determined, indicating that the choice of either HPS/DX or MH lamps should not affect growth response to UV-B radiation. This was true even though leaves of UV-B-irradiated plants grown under HPS/DX lamps have been shown to have greater chlorosis than those grown under MH lamps.