Effects of photoperiod and temperature on sugars and fructans in leaf blades, leaf sheaths and stems, and roots in relation to growth of Poa pratensis

Vegetative plants of Poa pratensis L. cv. Holt were cultivated in short days (SD; 8 h summer daylight) and in long days (LD; 8 h summer daylight + low intensity extension of 5 μmol m^-2 s^-1) at 12, 18 and 24°C in one experiment and at 9, 12, 15 and 18°C in another. Relative growth rate (RGR) as the mean of both experiments and all temperatures was 32% higher in LD than in SD between start of daylength treatment and first harvest, and 18% higher in LD than in SD between first and second harvest. Early in the daylength treatment period, more assimilates were allocated to storage in SD than in LD, so that at first harvest leaf sheaths and stems had 175% higher concentration of fructans in SD. Later this allocation pattern changed, and for the larger plants at the second harvest the differences in fructan concentrations were much smaller between the two daylengths. Both sugar and fructan concentrations were highest at low temperatures. The distribution of sugars and fructans varied from mostly sugars in the leaves to mostly fructans in leaf sheaths and stems and roots. The fructans were mainly high degree polymerization fructans. At least two series of fructans were present, and the dominant one was probably based upon kestose. It is concluded that allocation of assimilates to growth in leaf area instead of to storage may be important for the observed LD stimulation of dry matter production.     

Specific photoperiodic stimulation of dry matter production in a high-latitude cultivar of Poa pratensis

Vegetative plants of Poa pratensis L. cv. Holt (origin 69°N) raised in short days gave large and significant increases in plant dry weight, plant height and leaf area upon exposure to continuous light, compared with 8-h short days, at essentially identical daily inputs of radiant energy (8-h summer daylight ± low intensity extension). For example, by the fourth harvest (after 26, 34 and 46 days at 21, 15 and 9°C, respectively), the dry weights of plants in long days were 81, 163 and 195% greater than those of the corresponding short-day controls at the respective temperatures. Plant leaf areas in long days were between two and four times as large as control values by the end of the experiment. This was mainly due to increased leaf length caused by long-day stimulation of cell extension and division. However, the photoperiod did not affect the partitioning of assimilates amongst leaves, culms and stolons. Most of these effects could also be brought about by exogenous gibberellin application to plants in short days. However, in contrast to the effect of long days, gibberellin treatment also induced stem internode elongation even in these vegetative plants. Examination by standard growth analysis procedures revealed that the observed increases in relative growth rate were due primarily to increased net assimilation rate followed, several days later, by increases in leaf area ratio when newly-emerged leaves began to constitute a significant proportion of the leaf area. It is concluded that these reactions are of great adaptive significance for growth at the marginal temperatures prevailing at high latitudes.     

Gibberellins and the photoperiodic control of leaf growth in Poa pratensis

The role of gibberellins (GAs) in photoperiodic control of leaf elongation in Poa pratensis was studied by both application of exogenous GAs and analysis of endogenous GAs. Leaf elongation was strongly increased under long day (LD, 24 h) conditions at both 9 and 21°C, leaf length at 9°C LD being similar to that in plants grown in short days (SD, 8 h) at 21°C. However, even at 21°C leaf elongation was enhanced by LD. Exogenous GA₁ could completely compensate for LD at both 9 and 21°C. Gibberellins A₂₀, A₁₉ and A₄₄ could also partly replace LD, but they were significantly less active than GA₁, GA₅₃ was inactive when applied to plants grown at 9°C in SD and exhibited only marginal activity at 9°C LD and 21°C SD. The total level of GAs of the early 13-hydroxylation pathway (A₅₃, A₄₄, A₁₉, A₂₀ and A₁) increased rapidly when plants were transferred from SD to LD at 9°C. After transfer from 9 to 21°C, there was an increase in GA levels at both LD and SD, followed by a decrease under LD conditions. In all cases, GA₁₉ was the predominant GA, accounting for 60 to 80% of the analysed GAs. Levels of the bioactive GA₁ were low and increased transiently by LD four days after transfer from SD to LD. At both temperatures, the ratio GA₁₉ to GA₂₀ and GA₂₀ to GA₁ at 9°C was enhanced by LD compared with SD. Taken together, these results support the hypothesis that photoperiodic regulation of leaf elongation in Poa pratensis is GA-mediated, and they indicate a photoperiodic control of oxidation of GA₅₃ to GA₄₄ and GA₁₉ to GA₂₀, and perhaps also of 3β-hydroxylation of GA₂₀ to GA₁.     

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.     

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.     

Effects of exogenously applied growth regulators on shoot growth of inbred lines of Plantago major differing in relative growth rate: differential response to gibberellic acid and (2-chloroethyl)-trimethyl-ammonium chloride

The possibility of modulating shoot growth charaeteristics of seedlings of two inbred lines of Plantago major L., differing in relative growth rate (RGR), by exogenously applied 6-benzylaminopurine (BA), α-naphthalene acetic acid (NAA), (gibberellic acid (GA₃) and (2-chloroethyl)-trimethyl-ammonium chloride (CCC) was investigated. BA completely inhibited growth of the shoot at a concentration of 1 m M , while lower concentrations had no effect. NAA reduced growth of the shoot at 10 ü M , while 1 m M completely inhibited growth. Addition of 10 μ M GA₃ or higher stimulated shoot fresh weight up to 20% and leaf area up to 30% for the slow growing inbred line (W9), but less for the fast growing line (A4). Application of 1 m M CCC, an inhibitor of gibberellin metabolism, reduced growth of both inbred lines, but to a larger extent in the fast growing seedlings.
The lower shoot growth of W9 was associated with a lower specific leaf area (SLA) and a higher dry matter percentage of the shoot, as compared with A4. NAA reduced growth by reducing SLA and increasing leaf thickness, but the percentage dry matter of the leaves was unaffected. Stimulation of the shoot growth by GA₃ application was associated with higher SLA and lower dry matter percentage. Application of CCC had opposite effects on SLA and dry matter percentage as compared with GA₃. GA seems to be involved in the regulation of at least part of the genetic difference in RGR in Plantago major .     

Interaction of gibberellic acid and photoperiod on leaf sheath elongation in normal and gibberellin-insensitive genotype of wheat

Effect of gibberellic acid (GA₃) on leaf sheath elongation in a normal (cv. Møystad) and a gibberellin(GA)-insensitive (cv. Siete Cerros) genotype of wheat ( Triticum aestivum L.) were studied at 18 and 12°C under short (SD, 12 h) or long (LD, 24 h) photoperiod. Leaf sheath length in cv. Møystad was signficantly increased by exogenous GA₃ both under SD and LD. LD alone stimulated leaf sheath elongation and the combined effect of LD and GA₃ was additive, and there was no statistically signficant interaction between photoperiod and GA₃ concentrations. Leaf sheath length in cv. Siete Cerros was not significantly affected by GA₃ under any conditions. However, there was a highly significant stimulation of leaf sheath elongation by LD in cv. Siete Cerros as well. These results indicate that stimulation of elongation growth in wheat leaves by LD is not mediated by gibberellin.     

Inhibitory and promotive effects of gibberellic acid on floral initiation and development in Poa pratensis and Bromus inermis

Flowering in Poa pratensis L. cv. Holt and Bromus inermis Leyss. cv. Manchar requires exposure to short days (SD) for primary induction to occur, followed by long days (LD) to allow the inflorescence to develop. Weekly sprays with gibberellic acid (GA₃) during primary induction inhibited flower initiation in both P. pratensis and B. inermis . With 10⁻⁴ M GA₃ flowering of P. pratensis was suppressed even after an induction period of 10 weeks. Since both GA₃ and non-inductive LD conditions greatly stimulate leaf elongation, the degree of primary induction was closely negatively correlated with plant height (leaf sheath and blade length) at the end of the induction period. GA₃ application or the interpolation of LD during SD induction were most inhibitory during the later middle part of the SD period, whereas they were stimulatory near the beginning or immediately before the SD period. We suggest that changes in the portfolio or levels of endogenous gibberellins mediate photoperiodic control of growth and floral initiation in these plants. However, GA₃ sprays could not substitute for LD in causing heading and culm elongation in SD induced plants of the two species. The results are discussed in the light of results with other plants with dual floral induction requirements.     

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.     

Cell wall composition of leaves of an inherently fast- and an inherently slow-growing grass species

Differences in the relative growth rules of the inherently slow-growing Deschampsia flexuosa L. and the inherently fast-growing Holcus lanatus L. were reflected in cell wall synthesis in the elongation zone of the leaves. Leaf elongation rates depended on the size of the plant and ranged from 6 to 14 mm d^?1 in Deschampsia and from 12 to 42 mm d^?1 in Holcus. Anatomical data showed that the epidermis and vascular tissue are the important tissues controlling leaf extension. The cell wall polysaccharides of fully expanded leaves of the two species were identical in sugar composition. Enzymatic hydrolysis of polymeric sugars in the cell walls of the sheath and the lamina gave glucose (85%), arabinose (3.5%), fucose (0.5%), xylose (5.0%), mannose (0.5%), galaclose (0.8%) and galacturonic acid (3–4%). This composition applied throughout the blade and the sheath and did not change with ageing. Polysaccharides in the meristems of the two species showed identical sugar compositions with 51–55% glucose, 13–15% galactoronic acid and 13–14% arabinose as the main components. The extension zone was marked by a gradual increase of driselase-digestable polymers (per mm tissue) and a concurrent shift in sugar composition. The massive increase of glucose in the cell wall polymers of the elongation zone is probably caused by cellulose synthesis. The rate of synthesis of cell wall polysaccharides in Holcus was twice as high as that in Deschampsia. The slower-growing Deschampsia has more ferulic acid esterified with cell walls, which might contribute to the slowing of leaf growth. Lignin is not significantly deposited until growth has essentially ceased and is not responsible for the difference in growth rate.     

Differential selection of growth rate-related traits in wild barley, Hordeum spontaneum, in contrasting greenhouse nutrient environments

Across-species comparisons show that inherent variation in relative growth rate (RGR) and its underlying traits are correlated with habitat productivity. In this study, we test the hypothesis that growth rate-related traits confer differential selective effects in contrasting nutrient environments. We specifically test whether high RGR is targeted by selection in nutrient-rich environments whereas low values of traits that underlie RGR [specific leaf area (SLA), leaf mass fraction and leaf area ratio (LAR)] confer a direct fitness advantage in nutrient-poor environments, resulting in selection of low RGR as a correlated response. We measured RGR, its underlying component traits, and estimated fitness in a range of wild barley (Hordeum spontaneum) accessions grown under high and low nutrient conditions. Selection on component traits differed between the two environments, while total selection of RGR was not significant. Using multiple regression and path analysis to estimate direct fitness effects, a selective advantage of high LAR and SLA was demonstrated only under nutrient-rich conditions. While supporting the view that observed associations between habitat richness and some RGR-component traits reflect adaptation to differing nutrient regimes, our data suggest that direct selection targets component traits rather than RGR itself.     

刈割、施肥和浇水对垂穗披碱草补偿生长的影响

植物的补偿生长特性受放牧强度和生境资源获得性的影响。通过为期2年的野外控制实验,研究了刈割高度(留茬1cm、3cm及不刈割)、施肥(施、不施)和浇水(浇、不浇)处理对垂穗披碱草(Elymus natans)补偿生长的影响,并结合对各处理分株密度、比叶面积、净光合速率和相对生长率的变化研究,探讨了其补偿生长机制。结果表明:刈割后垂穗披碱草分株种群密度显著增加,补偿生长高度显著降低,比叶面积和相对生长率随刈割强度增加而呈上升趋势,叶片净光合速率变化不显著;施肥能显著增加垂穗披碱草的补偿生长高度、比叶面积、叶片净光合速率和相对生长率;浇水处理以及刈割、浇水、施肥处理之间的交互作用均不显著。可见,在刈割条件下,垂穗披碱草具有一定的密度补偿机制,但由于刈割抑制补偿性高生长,导致分株高度出现低补偿。因此,即使刈割后比叶面积和株高相对生长率显著增加,也不一定必然引起株高的超补偿;但施肥可显著提高垂穗披碱草的补偿能力,增加耐牧性,证实了改进后的限制资源模型的预测。     

Ameliorating effect of triacontanol on salt stressederythrina variegata seedlings. Changes in growth,biomass, pigments and solute accumulation

Erythrina variegata Lam. seedlings were grown under low (100 mM NaCl) and high (250 mM NaCl) salinity. Seedlings exposed to high salinity for 10 d showed significant reduction in growth rate and biomass production while the root/shoot ratio increased. In contrast to pigment and protein contents, starch and saccharide contents increased in salt stressed seedlings. When the seedlings were subsequently sprayed with triacontanol (1 mg kg^-1) the salinity effect was partially ameliorated and growth, biomass, chlorophyll and carotenoid contents increased.     

Variation in the components of relative growth rate in 10 Acacia species from contrasting environments

In this study we assessed the inherent relative growth rate (RGR) under controlled environment conditions of 10 contrasting Acacia species from semi-arid and mesic environments. For several of the species, compound pinnate leaves produced early in the seedling stage, were gradually replaced by phyllodes (expanded petioles that form simple lamina). Other species either did not form phyllodes, or only did so to a minor degree by the end of the study. Phyllode production was dominant in the four slow-growing Acacia species from semi-arid environments (A. aneura, A. colei, A. coriacea and A. tetragonophylla), with leaf production being exclusive or dominant in five (A. dealbata, A. implexa, A. mearnsii, A. melanoxylon and A. irrorata) of the six faster-growing species from mesic environments. The exception was A. saligna which was fast growing but did produce phyllodes. From a carbon economy perspective, slow growth in the semi-arid species was not associated with lower net assimilation rates or less plant mass allocated to foliage. Rather, the primary factor associated with their slow growth was a smaller foliage area per unit foliage mass. This was true for comparisons based on the mean over all harvests or at set plant masses. The production of phyllodes by the semi-arid species substantially reduced foliage area per unit foliage mass, as this was lower for phyllodes than leaves in all species. To assess the impact that phyllode production had on ontogenetic changes in RGR, we modelled the situation where only leaves were formed. This analysis showed that changing from leaves to phyllodes substantially reduced the RGR. There was little difference in plant nitrogen concentration or the ratio of foliage nitrogen to plant nitrogen between the species. This resulted in foliage nitrogen productivity (dry mass gain per unit foliage nitrogen and time) being directly proportional to foliage area per unit foliage mass between species. We concluded that a smaller foliage area per unit foliage mass and phyllode production are the primary factors associated with lower RGR in contrasting Acacia species.     

The influence of gibberellic acid and sulfur fertilization rate on growth and S-use efficiency of mustard (<Emphasis Type="Italic">Brassica juncea</Emphasis>)

Earlier research has shown that exogenous gibberellic acid (GA₃) application increases shoot growth, photosynthesis and soil nitrogen (N) utilisation in mustard (Brassica juncea L. Czern & Coss.). Mustard has a high sulfur (S) requirement. Its assimilatory pathway is well coordinated with N and dependent on photosynthesis. Thus, the higher photosynthate production and an efficient use of N with the use of GA₃ could result in an increase in S-use efficiency of the crop. The research was, therefore, carried out to study the effects of 10~M GA₃ spray on specific leaf area, plant dry mass, leaf carbon dioxide exchange rate (CER), plant growth rate (PGR), relative growth rate (RGR), net assimilation rate (NAR) and S-use efficiency (SUE) of mustard treated with 0, 100 or 200 mg S kg^–1 soil levels. Plants treated with 100~mg S kg^–1 soil and receiving GA₃ treatment showed increased specific leaf area and dry mass accumulation compared to the control. At 0~mg S kg^–1 soil, N and S concentrations were reduced. They increased with increasing S supply. GA₃ application significantly increased N and S concentrations further. A two-fold increase in SUE in GA₃-treated plants at 100~mg S kg^–1 soil was noted in comparison to the control. SUE was not increased under excess S conditions beyond 100~mg S kg^–1 soil. The increase in SUE was through increase in the growth, CER and use efficiency of N by the crop due to GA₃ application.