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.     

Thermal acclimation of leaf and root respiration: an investigation comparing inherently fast- and slow-growing plant species

We investigated the extent to which leaf and root respiration (R) differ in their response to short‐ and long‐term changes in temperature in several contrasting plant species (herbs, grasses, shrubs and trees) that differ in inherent relative growth rate (RGR, increase in mass per unit starting mass and time). Two experiments were conducted using hydroponically grown plants. In the long‐term (LT) acclimation experiment, 16 species were grown at constant 18, 23 and 28 °C. In the short‐term (ST) acclimation experiment, 9 of those species were grown at 25/20 °C (day/night) and then shifted to a 15/10 °C for 7 days. Short‐term Q₁₀ values (proportional change in R per 10 °C) and the degree of acclimation to longer‐term changes in temperature were compared. The effect of growth temperature on root and leaf soluble sugar and nitrogen concentrations was examined. Light‐saturated photosynthesis (A_sat) was also measured in the LT acclimation experiment. Our results show that Q₁₀ values and the degree of acclimation are highly variable amongst species and that roots exhibit lower Q₁₀ values than leaves over the 15–25 °C measurement temperature range. Differences in RGR or concentrations of soluble sugars/nitrogen could not account for the inter‐specific differences in the Q₁₀ or degree of acclimation. There were no systematic differences in the ability of roots and leaves to acclimate when plants developed under contrasting temperatures (LT acclimation). However, acclimation was greater in both leaves and roots that developed at the growth temperature (LT acclimation) than in pre‐existing leaves and roots shifted from one temperature to another (ST acclimation). The balance between leaf R and A_sat was maintained in plants grown at different temperatures, regardless of their inherent relative growth rate. We conclude that there is tight coupling between the respiratory acclimation and the temperature under which leaves and roots developed and that acclimation plays an important role in determining the relationship between respiration and photosynthesis.     

Effects of N-supply on the rates of photosynthesis and shoot and root respiration of inherently fast- and slow-growing monocotyledonous species

Are there intrinsic differences in the rates of photosynthesis, shoot- and root-respiration between inherently fast- and slow-growing monocotyledons at high and low nitrogen supply? To analyze this question we grew 5 monocotyledons, widely differing in their inherent relative growth rate at high and low nitrogen supply in a growth room. Nitrate was exponentially added to the plants, enabling us to compare inherent differences in plant characteristics, without any effect of species differences in the ability to take up nutrients. At high nitrogen supply, the fast-growing species from productive habitats had a higher photosynthetic nitrogen use efficiency and rate of root respiration than the slow-growing ones from unproductive habitats. Only minor differences were observed in their rates of photosynthesis and shoot respiration per unit leaf area. At low nitrogen supply, the rates of photosynthesis and shoot- and root respiration decreased for all species, even though there were no longer any differences in these processes between inherently fast- and slow-growing species. The photosynthetic nitrogen use efficiency increased for all species, and no differences were found among species. Differences in the photosynthetic nitrogen use efficiency among species and nitrogen treatments are discussed in terms of the utilization of the photosynthetic apparatus, whereas differences in respiration rate are discussed in terms of the energy demand for growth, maintenance and ion uptake and their related specific respiratory energy costs. It is concluded that the relatively high abundance of slow-growing species compared to fast-growing ones in unproductive habitats is unlikely to be explained by differences in rates of photosynthesis and respiration or in photosynthetic nitrogen use efficiency.     

The chemical composition and anatomical structure of leaves of grass species differing in relative growth rate

To arrive at a better understanding of variation in specific leaf mass (SLM, leaf weight per unit leaf area), we investigated the chemical composition and anatomical structure of the leaves of 14 grass species varying in potential relative growth rate. Expressed on a dry weight basis, the fast-growing grass species with low SLM contained relatively more minerals and organic N-compounds, whereas slow-growing species with high SLM contained more (hemi)cellulose and lignin. However, when expressed per unit leaf area, organic N-compounds, (hemi)cellulose, total structural carbohydrates and organic acids increased with increasing SLM. For the 14 grasses, no trend with SLM was found for the leaf volume per unit leaf area. Leaf density was positively correlated with SLM. Variation in density was not caused by variation in the proportion of intercellular spaces. The proportion of the total volume occupied by mesophyll and veins did not differ either. A high SLM was caused, at least partly, by a high proportion of non-veinal sclerenchymatic cells per cross-section. The epidermal cell area was negatively correlated with SLM. We conclude that the differences in SLM and in the relative growth rate (RGR) between fast- and slow-growing grass species are based partly on variation in anatomical differentiation and partly on chemical differences within cell types.     

Root and leaf attributes accounting for the performance of fast- and slow-growing grasses at different nutrient supply

Despite their difference in potential growth rate, the slow-growing Brachypodium pinnatum and the fast-growing Dactylis glomerata co-occur in many nutrient-poor calcareous grasslands. They are known to respond differently to increasing levels of N and P. An experiment was designed to measure which characteristics are affected by nutrient supply and contribute to the ecological performance of these species. Nutrient acquisition and root and shoot traits of these grasses were studied in a garden experiment with nine nutrient treatments in a factorial design of 3 N and 3 P levels each. D. glomerata was superior to B. pinnatum in nutrient acquisition and growth in all treatments. B. pinnatum was especially poor in P acquisition. Both species responded to increasing N supply and to a lesser extent to increasing P supply by decreasing their root length and increasing their leaf area per total plant weight. D. glomerata showed a higher plasticity. In most treatments, the root length ratio (RLR) and the leaf area ratio (LAR) were higher for D. glomerata. A factorization of these parameters into components expressing biomass allocation, form (root fineness or leaf thickness) and density (dry matter content) shows that the low density of the biomass of D. glomerata was the main cause for the higher RLR and LAR. The biomass allocation to the roots showed a considerable plasticity but did not differ between the species. B. pinnatum had the highest leaf weight ratio. Root fineness was highly plastic in D. glomerata, the difference with B. pinnatum being mainly due to the thick roots of D. glomerata at high nutrient supply. The leaf area/leaf fresh weight ratio did not show any plasticity and was slightly higher for B. pinnatum. It is concluded, that the low density of the biomass of D. glomerata is the pivotal trait responsible for its faster growth at all nutrient levels. It enables simultaneously a good nutrient acquisition capacity by the roots as well as a superior carbon acquisition by the leaves. The high biomass density of B. pinnatum will then result in a lower nutrient requirement due to a slower turnover, which in the long term is advantageous under nutrient-poor conditions.     

XET-related genes and growth kinematics in barley leaves

DV, displacement velocity
GA₃, gibberellic acid
REGR, relative elemental growth rate
XET, xyloglucan-endotransglycosylase

Recently Schünmann et al. (1997 ; Plant, Cell and Environment 20, 1439–1450) investigated the correlation of spatial patterns of xyloglucan-endotransglycosylase (XET) activity, XET-related mRNAs, and growth in elongating barley ( Hordeum vulgare L.) leaves. Here, methodological difficulties in the kinematic growth analysis are discussed, and it is concluded that the role that XET-related gene activity plays in the control of spatial growth patterns remains undetermined.     

A functional-structural model of elongation of the grass leaf and its relationships with the phyllochron

The emergence of a regular phyllochron from the dynamic processes of leaf initiation, leaf elongation and whorl construction suggests causal relationships between leaf elongation and leaf emergence. This paper presents a hypothesis as to how the ontogeny of the growth zone of leaves is triggered by emergence events, and implements it in a dynamic model of leaf elongation. Two different experiments, presenting two contrasted cases of relationships between leaf emergence and kinetics of leaf elongation, were analysed and interpreted with the model in terms of the functioning of the growth zone. Analysis of elongation kinetics revealed that the hypothesis allows for several contrasted elongation patterns that were observed, and for a regular phyllochron emerging from the variable dynamic of elongation. The model was able to simulate these patterns, and helped to identify the mechanisms underlying the key points of the analysis. The hypothesis is not demonstrated, but its coherence and robustness are established, which should inform a renewal of the modelling of leaf elongation in architectural models.     

Biophysical limitation of cell elongation in cereal leaves

Grass leaves grow from the base. Unlike those of dicotyledonous plants, cells of grass leaves expand enclosed by sheaths of older leaves, where there is little or no transpiration, and go through developmental stages in a strictly linear arrangement. The environmental or developmental factor that limits leaf cell expansion must do so through biophysical means at the cellular level: wall-yielding, water uptake and solute supply are all candidates. This Botanical Briefing looks at the possibility that tissue hydraulic conductance limits cell expansion and leaf growth. A model is presented that relates pathways of water movement in the elongation zone of grass leaves to driving forces for water movement and to anatomical features. The bundle sheath is considered as a crucial control point. The relative importance of these pathways for the regulation of leaf growth and for the partitioning of water between expansion and transpiration is discussed.     

Phosphorus nutrition and mycorrhiza effects on grass leaf growth. P status- and size-mediated effects on growth zone kinematics

This study tested whether leaf elongation rate (LER, mm h(-1)) and its components--average relative elemental growth rate (REGRavg, mm mm(-1) h(-1)) and leaf growth zone length (L(LGZ), mm)--are related to phosphorus (P) concentration in the growth zone (P(LGZ) mg P g(-1) tissue water) of Lolium perenne L. cv. Condesa and whether such relationships are modified by the arbuscular mycorrhizal fungus (AMF) Glomus hoi. Mycorrhizal and non-mycorrhizal plants were grown at a range of P supply rates and analysed at either the same plant age or the same tiller size (defined by the length of the sheath of the youngest fully expanded leaf). Both improved P supply (up to 95%) and AMF (up to 21%) strongly increased LER. In tillers of even-aged plants, this was due to increased REGRavg and L(LGZ). In even-sized tillers, it was exclusively due to increased REGRavg. REGRavg was strictly related to P(LGZ) (r2 = 0.95) and independent of tiller size. Conversely, L(LGZ) strictly depended on tiller size (r2 = 0.88) and not on P(LGZ). Hence, P status affected leaf growth directly only through effects on relative tissue expansion rates. Symbiosis with AMF did not modify these relationships. Thus, no evidence for P status-independent effects of AMF on LER was found.     

Effects of nitrogen supply on the anatomy and chemical composition of leaves of four grass species belonging to the genus Poa, as determined by image-processing analysis and pyrolysis–mass spectrometry

Previous experiments have shown that the anatomy and chemical composition of leaves of inherently fast- and slow-growing grass species, grown at non-limiting nitrogen supply, differ systematically. The present experiment was carried out to investigate whether these differences persist when the plants are grown at an intermediate or a very low nitrogen supply. To this end, the inherently fast-growing Poa annua L. and Poa trivialis L., and the inherently slow-growing Poa compressa L. and Poa pratensis (L.) Schreb. were grown hydroponically at three levels of nitrate supply: at optimum (RGR_max) and at relative addition rates of 100 and 50 mmol N (mol N)^?1 d^?1 (RAR₁₀₀ and RAR₅₀), respectively. As expected, at the lowest N supply, the potentially fast-growing species grew at the same rate as the inherently slow-growing ones. Similarly, the differences in leaf area ratio (LAR, leaf area:total dry mass), specific leaf area (SLA, leaf arear:leaf dry mass) and leaf mass ratio (LMR, leaf dry mass:total dry mass) disappeared. Under optimal conditions, the fast-growing species differed from the slow-growing ones in that they had a higher N concentration. There were no significant differences in C concentration. With decreasing N supply, the total N concentration decreased and the differences between the species disappeared. The total C concentration increased for the fast-growing species and decreased for the slow-growing ones, i.e. the small, but insignificant, difference in C concentration between the species at RGR_max increased with decreasing N supply. The chemical composition of the leaves at low N supply, analysed in more detail by pyrolysis–mass spectrometry, showed an increase in the relative amounts of guaiacyl lignin, cellulose and hemicellulose, whereas those of syringyl lignin and protein decreased. The anatomy and morphology of the leaves of the four grass species differing in RGR_max were analysed by image-processing analysis. The proportion of the total volume occupied by mesophyll plus intercellular spaces and epidermis did not correlate with the amount of leaf mass per unit leaf area (specific leaf mass, SLM) at different N supply. The higher SLM at low N supply was caused partly by a high proportion of non-veinal sclerenchymatic cells per cross-section and partly by the smaller volume of epidermal cells. We conclude that the decrease in relative growth rate (and increase in SLM) at decreasing N supply is partly due to chemical and anatomical changes. The differences between the fast- and slow-growing grass species at an optimum nutrient supply diminished when plants were growing at a limiting nitrogen supply.     

The decrease in growth of phosphorus-deficient maize leaves is related to a lower cell production

The spatial distribution of leaf elongation and adaxial epidermal cell production in leaf 6 of maize (Zea mays L. cv. Cecilia) plants grown in a growth chamber under two contrasting availabilities of P in the soil was investigated. Lower displacement velocities from 32.5 mm from leaf base and a shorter growth zone were found in low P (LP) leaves compared with control leaves. P deficiency significantly diminished maximum relative elemental growth rate and shifted its location closer to the leaf base. Cells were significantly longer in LP than in control leaves for all positions from the leaf base except at the end of the growth zone. For both treatments it took a similar time for a cell situated at the leaf base to reach the limit of the growth zone. The average length of the cell division zone was decreased by 21% in LP leaves. Significant differences were found in cell production and cell division rates from 12.5 mm from the leaf base although maximum values were similar between P treatments. A shorter zone of cell division with lower cell production rates along most of its length was the regulatory event that decreased cell production, and ultimately leaf elongation rates, in P‐deficient maize plants.     

The contribution of roots and shoots to whole plant nitrate reduction in fast- and slow-growing grass species

The hypothesis was tested that slow-growing grass species perform a greater proportion of total plant NO3- reduction in their roots than do fast-growing grasses. Eight grass species were selected that differed in maximum relative growth rate (RGR) and net NO3- uptake rate (NNUR). Plants were grown with free access to nutrients in hydroponics under controlled-environment conditions. The site of in vivo NO3- reduction was assessed by combining in vivo NO3- reductase activity (NRA) assays with biomass allocation data, and by analysing the NO3- to amino acid ratio of xylem sap. In vivo NRA of roots and shoots increased significantly with increasing NNUR and RGR. The proportion of total plant NO3- reduction that occurs in roots was found to be independent of RGR and NNUR, with the shoot being the predominant site of NO3- reduction in all species. The theoretical maximum proportion of whole plant nitrogen assimilation that could take place in the roots was calculated using information on root respiration rates, RGR, NNUR, and specific respiratory costs associated with growth, maintenance and ion uptake. The calculated maximum proportion that the roots can contribute to total plant NO3- reduction was 0.37 and 0.23 for the fast-growing Dactylis glomerata L. and the slow-growing Festuca ovina L., respectively. These results indicate that slow-growing grass species perform a similar proportion of total plant NO3- reduction in their roots to that exhibited by fast-growing grasses. Shoots appear to be the predominant site of whole plant NO3- reduction in both fast- and slow-growing grasses when plants are grown with free access to nutrients.     

Cell wall structure and biogenesis in Aspergillus species

Aspergillus species are among the most important filamentous fungi from the viewpoints of industry, pathogenesis, and mycotoxin production. Fungal cells are exposed to a variety of environmental stimuli, including changes in osmolality, temperature, and pH, which create stresses that primarily act on fungal cell walls. In addition, fungal cell walls are the first interactions with host cells in either human or plants. Thus, understanding cell wall structure and the mechanism of their biogenesis is important for the industrial, medical, and agricultural fields. Here, we provide a systematic review of fungal cell wall structure and recent findings regarding the cell wall integrity signaling pathways in aspergilli. This accumulated knowledge will be useful for understanding and improving the use of industrial aspergilli fermentation processes as well as treatments for some fungal infections.     

The response of fast- and slow-growing Acacia species to elevated atmospheric CO2: an analysis of the underlying components of relative growth rate

In this study we assessed the impact of elevated CO₂ with unlimited water and complete nutrient on the growth and nitrogen economy of ten woody Acacia species that differ in relative growth rate (RGR). Specifically, we asked whether fast- and slow-growing species systematically differ in their response to elevated CO₂. Four slow-growing species from semi-arid environments (Acacia aneura, A. colei, A. coriacea and A. tetragonophylla) and six fast-growing species from mesic environments (Acacia dealbata, A. implexa, A. mearnsii, A. melanoxylon, A. irrorata and A. saligna) were grown in glasshouses with either ambient (˜350 ppm) or elevated (˜700 ppm) atmospheric CO₂. All species reached greater final plant mass with the exception of A. aneura, and RGR, averaged across all species, increased by 10% over a 12-week period when plants were exposed to elevated CO₂. The stimulation of RGR was evident throughout the 12-week growth period. Elevated CO₂ resulted in less foliage area per unit foliage dry mass, which was mainly the result of an increase in foliage thickness with a smaller contribution from greater dry matter content per unit fresh mass. The net assimilation rate (NAR, increase in plant mass per unit foliage area and time) of the plants grown at elevated CO₂ was higher in all species (on average 30% higher than plants in ambient CO₂) and was responsible for the increase in RGR. The higher NAR was associated with a substantial increase in foliar nitrogen productivity in all ten Acacia species. Plant nitrogen concentration was unaltered by growth at elevated CO₂ for the slow-growing Acacia species, but declined by 10% for faster-growing species. The rate of nitrogen uptake per unit root mass was higher in seven of the species when grown under elevated CO₂, and leaf area per unit root mass was reduced by elevated CO₂ in seven of the species. The absolute increase in RGR due to growth under elevated CO₂ was greater for fast- than for slow-growing Acacia species. Received: 21 December 1998 / Accepted: 31 May 1999     

Control of Leaf Growth and its Role in Determining Variation in Plant Growth Rate from an Ecological Perspective

Abstract: Plants vary widely in their relative growth rate (RGR), be it dependent on environmental conditions or due to their genetic background. In a comparison of the RGR of grasses growing under different environmental conditions, variation in RGR tends to correlate with that in the leaf elongation rate (LER). When different species or genotypes thereof are compared under identical growing conditions, variation in LER may or may not correlate with that in RGR, depending on the comparison. However, since RGR is described by an exponential equation, whereas LER is mainly a linear process, we conclude that any correlation between RGR and LER must be fortuitous. That is, exponential growth must be due to increases with time in plant traits such as 1) leaf dry mass per unit leaf length invested per unit time, and/or 2), i.e., the total LER of all the growing leaves at one point in time. The latter can be achieved as follows: 1) each subsequent leaf has a higher LER than the preceding one; 2) leaves appear at an increasing rate; 3) the duration of the process of leaf elongation increases for subsequent leaves. In this review, we only explore possible factors that account for changes in with time, in different genotypes and under different environmental conditions. Inherent variation in LER of individual leaves and variation due to environmental factors may reflect variation in the rate of cell division and/or in cell elongation.     

From individual leaf elongation to whole shoot leaf area expansion: a comparison of three Aegilops and two Triticum species

BACKGROUND AND AIMS: Rapid leaf area expansion is a desirable trait in the early growth stages of cereal crops grown in low-rainfall areas. In this study, the traits associated with inherent variation in early leaf area expansion rates have been investigated in two wheat species (Triticum aestivum and T. durum) and three of its wild relatives (Aegilops umbellulata, A. caudata and A. tauschii) to find out whether the Aegilops species have a faster leaf area expansion in their early developmental stage than some of the current wheat species. METHODS: Growth of individual leaves, biomass allocation, and gas exchange were measured on hydroponically grown plants for 4 weeks. KEY RESULTS: Leaf elongation rate (LER) was strongly and positively correlated with leaf width but not with leaf elongation duration (LED). The species with more rapidly elongating leaves showed a faster increase with leaf position in LER, leaf width and leaf area, higher relative leaf area expansion rates, and more biomass allocation to leaf sheaths and less to roots. No differences in leaf appearance rate were found amongst the species. CONCLUSIONS: Aegilops tauschii was the only wild species with rapid leaf expansion rates similar to those of wheat, and it achieved the highest photosynthetic rates, making it an interesting species for further study.     

Measurement of a growth-induced water potential gradient in tall fescue leaves

Spatial distribution of cell turgor pressure, cell osmotic pressure and relative elemental growth rate were measured in growing tall fescue leaves ( Festuca arundinacea ). Cell turgor pressure (measured with a pressure probe) was c . 0.55 MPa in expanding cells but increased steeply (+0.3 MPa) in cells where elongation had stopped. However, cell osmotic pressure (measured with a picolitre osmometer) was almost constant at 0.85 MPa throughout the leaf. The water potential difference between the growth zone and the mature zone (0.3 MPa) was interpreted as a growth-induced water potential gradient. This and further implications for the mechanism of growth control are discussed.     

Dynamics of intracellular mannan and cell wall folding in the drought responses of succulent Aloe species

Plants have evolved a multitude of adaptations to survive extreme conditions. Succulent plants have the capacity to tolerate periodically dry environments, due to their ability to retain water in a specialized tissue, termed hydrenchyma. Cell wall polysaccharides are important components of water storage in hydrenchyma cells. However, the role of the cell wall and its polysaccharide composition in relation to drought resistance of succulent plants are unknown. We investigate the drought response of leaf‐succulent Aloe (Asphodelaceae) species using a combination of histological microscopy, quantification of water content, and comprehensive microarray polymer profiling. We observed a previously unreported mode of polysaccharide and cell wall structural dynamics triggered by water shortage. Microscopical analysis of the hydrenchyma cell walls revealed highly regular folding patterns indicative of predetermined cell wall mechanics in the remobilization of stored water and the possible role of homogalacturonan in this process. The in situ distribution of mannans in distinct intracellular compartments during drought, for storage, and apparent upregulation of pectins, imparting flexibility to the cell wall, facilitate elaborate cell wall folding during drought stress. We conclude that cell wall polysaccharide composition plays an important role in water storage and drought response in Aloe.     

Interaction of photoperiod and gibberellin on growth and photosynthesis of high-latitude Poa pratensis

The relative growth rate of pot-grown plants of Poa pratensis L. cv. Holt, origin 69s°N, was increased by 20–40% by photoperiod extension with low intensity incandescent light from 8 to 24 h at 9–21°C. The main increase occurred over the 14 to 18 h photoperiod range. The true photoperiodic nature of the response was demonstrated by the effectiveness of night interruption in stimulating growth. Fortnightly sprayings with gibberellic acid (GA₃) (3 × 10^-6 to 3 × 10^-5 M ) mimicked all the effects of long days, whereas (2-chloroethyl)-trimethylammonium chloride (CCC) counteracted the effects of long days. Both growth substances exhibited pronounced interactions with photoperiod, GA₃ being most effective in short days and CCC in long days. The growth stimulation, whether caused by long days or GA₃, was exerted mainly through increases in individual and total leaf area. This was associated with a reduction in CO₂, exchange rate and a parallel fall in specific leaf weight. Proportionally, however, the increase in leaf area was greater than the fall in CO₂ exchange rate, resulting in a 38 to 118% increase in photosynthesis per leaf. No evidence was found of any direct and promotive effect of transition to long days on the CO₂ exchange rate of already expanded leaves.