Effect of nitrogen supply on leaf appearance, leaf growth, leaf nitrogen economy and photosynthetic capacity in maize (Zea mays L.)

Leaf area growth and nitrogen concentration per unit leaf area, N-a (g m(-2) N) are two options plants can use to adapt to nitrogen limitation. Previous work indicated that potato (Solanum tuberosum L.) adapts the size of leaves to maintain Na and photosynthetic capacity per unit leaf area. This paper reports on the effect of N limitation on leaf area production and photosynthetic capacity in maize, a C4 cereal. Maize was grown in two experiments in pots in glasshouses with three (0.84-6.0 g N pot(-1)) and five rates (0.5-6.0 g pot(-1)) of N. Leaf tip and ligule appearance were monitored and final individual leaf area was determined. Changes with leaf age in leaf area, leaf N content and light-saturated photosynthetic capacity, P a,, were measured on two leaves per plant in each experiment. The final area of the largest leaf and total plant leaf area differed by 16 and 29% from the lowest to highest N supply, but leaf appearance rate and the duration of leaf expansion were unaffected. The N concentration of expanding leaves (N-a or %N in dry matter) differed by at least a factor 2 from the lowest to highest N supply. A hyperbolic function described the relation between P-max and N-a. The results confirm the 'maize strategy': leaf N content, photosynthetic capacity, and ultimately radiation use efficiency is more sensitive to nitrogen limitation than are leaf area expansion and light interception. The generality of the findings is discussed and it is suggested that at canopy level species showing the 'potato strategy' can be recognized from little effect of nitrogen supply on radiation use efficiency, while the reverse is true for species showing the 'maize strategy' for adaptation to N limitation. (c) 2004 Elsevier B.V. All rights reserved.

Effects of nitrogen supply on xylem cytokinin delivery, transpiration and leaf expansion of pea genotypes differing in xylem-cytokinin concentration

The rms2 and rms4 pea ( Pisum sativum L.) branching mutants have higher and lower xylem-cytokinin concentration, respectively, relative to wild type (WT) plants. These genotypes were grown at two levels of nitrogen (N) supply for 18 - 20 d to determine whether or not xylem-cytokinin concentration (X-CK) or delivery altered the transpiration and leaf growth responses to N deprivation. Xylem sap was collected by pressurising de-topped root systems. As sap-flow rate increased, X-CK declined in WT and rms2, but did not change in rms4. When grown at 5.0 mM N, X-CKs of rms2 and rms4 were 36% higher and 6-fold lower, respectively, than WT at sap-flow rates equivalent to whole-plant transpiration. Photoperiod cytokinin (CK) delivery rates ( the product of transpiration and X-CK) decreased more than 6-fold in rms4. Growth of plants at 0.5 mM N had negligible (< 10%) effects on transpiration rates expressed on a leaf area basis in WT and rms4, but decreased transpiration rates of rms2. The low-N treatment decreased leaf expansion by 20 - 25% and expanding leaflet N concentration by 15%. These changes were similar in all genotypes. At sap-flow rates equivalent to whole-plant transpiration, the low N treatment decreased X-CK in rms2 but had no discernible effect in WT and rms4. Since the low N treatment decreased transpiration of all genotypes, photoperiod CK delivery rates also decreased in all genotypes. The similar leaf growth response of all genotypes to N deprivation despite differences in both absolute and relative X-CKs and deliveries suggests that shoot N status is more important in regulating leaf expansion than xylem-supplied cytokinins. The decreased X-CK and transpiration rate of rms2 following N deprivation suggests that changes in xylem-supplied CKs may modify water use.

Adaptive responses of wild mungbean (Vigna radiata ssp sublobata) to photo-thermal environment. II. Growth, biomass, and seed yield

The leaf growth, dry matter production, and seed yield of 11 wild mungbean ( Vigna radiata ssp. sublobata) accessions of diverse geographic origin were observed under natural and artificial photoperiod temperature conditions, to determine the extent to which genotypic differences could be attributed to adaptive responses to photo-thermal environment. Environments included serial sowings in the field in SE Queensland, complemented by artificial photoperiod extension and controlled-environment growth rooms. Photo-thermal environment influenced leaf growth, total dry matter production ( TDM), and seed yield directly, through effects of ( mainly cool) temperature on growth, and indirectly, through effects on phenology. In terms of direct effects, leaf production, leaf expansion, and leaf area were all sensitive to temperature, with implied base temperatures higher than usually observed in cultivated mungbean ( V. radiata ssp. radiata). Genotypic sensitivity to temperature varied systematically with accession provenance and appeared to be of adaptive significance. In terms of the indirect effects of photo-thermal environment, genotypic and environmental effects on TDM were positively related to changes in total growth duration, and harvest index was negatively related to the period from sowing to flowering, similar to cultivated mungbean. However, seed yield was positively related to the duration of reproductive growth, reflecting the indeterminate growth habit of the wild accessions. As a consequence, the wild accessions are more responsive to favourable environments than typically observed in cultivated mungbean, which is determinate in habit. It is suggested that the introduction of the indeterminate trait into mungbean from the wild subspecies would increase the responsiveness of mungbean to favourable environments, analogous to that of black gram ( V. mungo). Although the wild subspecies appeared more sensitive to cool temperature than cultivated mungbean, it may provide a source of tolerance to the warmer temperatures experienced during the wet season in the tropics.

Summer dormancy in perennial temperate grasses

Background and Aims Dormancy has been extensively studied in plants which experience severe winter conditions but much less so in perennial herbaceous plants that must survive summer drought. This paper reviews the current knowledge on summer dormancy in both native and cultivated perennial temperate grasses originating from the Mediterranean Basin, and presents a unified terminology to describe this trait. Scope Under severe drought, it is difficult to separate the responses by which plants avoid and tolerate dehydration from those associated with the expression of summer dormancy. Consequently, this type of endogenous (endo-) dormancy can be tested only in plants that are not subjected to moisture deficit. Summer dormancy can be defined by four criteria, one of which is considered optional: (1) reduction or cessation of leaf production and expansion; (2) senescence of mature foliage; (3) dehydration of surviving organs; and (4, optional) formation of resting organs. The proposed terminology recognizes two levels of summer dormancy: (a) complete dormancy, when cessation of growth is associated with full senescence of foliage and induced dehydration of leaf bases; and (b) incomplete dormancy, when leaf growth is partially inhibited and is associated with moderate levels of foliage senescence. Summer dormancy is expressed under increasing photoperiod and temperature. It is under hormonal control and usually associated with flowering and a reduction in metabolic activity in meristematic tissues. Dehydration tolerance and dormancy are independent phenomena and differ from the adaptations of resurrection plants. Conclusions Summer dormancy has been correlated with superior survival after severe and repeated summer drought in a large range of perennial grasses. In the face of increasing aridity, this trait could be used in the development of cultivars that are able to meet agronomic and environmental goals. It is therefore important to have a better understanding of the genetic and environmental control of summer dormancy.

Xylem-borne cytokinins: still in search of a role?

Leaf expansion and xylem cytokinin concentration ([X-CK]) decrease in response to nitrogen (N) deprivation. Debate continues over cause, effect, and correlation. Supporting studies provide, at best, correlative evidence that [X-CK] controls leaf growth in response to N-deprivation, while dissenting studies indicate that leaf growth responses to N can be independent of changes in X-CK supply to leaves. A model is proposed to evaluate the physiological significance to leaf growth of changes in plant and environment N concentrations, and plant CK concentrations.

Effects of boron foliar applications on vegetative and reproductive growth of sunflower

Foliar application may be used to supply boron (B) to a crop when B demands are higher than can be supplied via the soil. While B foliar sprays have been used to correct B deficiency in sunflower (Helianthus annuus L.) in the field, no studies have determined the amount of B taken up by sunflower plant parts via foliar application. A study was conducted in which sunflower plants were grown at constant B concentration in nutrient solution with adequate B (46 mum) or with limited B supply (0.24, 0.40 and 1.72 mum) using Amberlite IRA-743 resin to control B supply. At the late vegetative stage of growth (25 and 35 d after transplanting), two foliar sprays were applied of soluble sodium tetraborate (20.8 % B) each at 0, 28, 65, 120 and 1200 mm (each spray equivalent to 0, 0.03, 0.07, 0.13 and 1.3 kg B ha(-1) in 100 L water ha(-1)). The highest rate of B foliar fertilization resulted in leaf burn but had no other evident detrimental effect on plant growth. Under B-deficient conditions, B foliar application increased the vegetative and reproductive dry mass of plants. Foliar application of 28-1200 mm B increased the total dry mass of the most B-deficient plants by more than three-fold and that of plants grown initially with 1.72 mum B in solution by 37-49 %. In this latter treatment, the dry mass of the capitulum was similar to that achieved under control conditions, but in no instance was total plant dry mass similar to that of the control. All B foliar spray rates increased the B concentration in various parts of the plant tops, including those that developed after the sprays were applied, but the B concentration in the roots was not increased by B foliar application. The B concentration in the capitulum of the plants sprayed at the highest rate was between 37 and 93 % of that in the control plants. This study showed that B foliar application was of benefit to B-deficient sunflower plants, increasing the B status of plant tops, including that of the capitulum which developed after the B sprays were applied. (C) 2003 Annals of Botany Company.

The effect of timing and severity of water deficit on growth, development, yield accumulation and nitrogen fixation of mungbean

Mungbean (Vigna radiata L.), as a dryland grain legume, is exposed to varying timing and severity of water deficit, which results in variability in grain yield, nitrogen accumulation and grain quality. In this field study, mungbean crops were exposed to varying timing and severity of water deficit in order to examine: (1) contribution of the second flush of pods to final grain yield with variable timing of relief from water deficit, (2) the sensitivity to water deficit of the accumulation of biomass and nitrogen (N) and its partitioning to grain, and (3) how the timing of water deficit affects the pattern of harvest index (HI) increase through pod filling. The results showed that the contribution of the second flush to final yield is highly variable (1-56%) and can be considerable, especially where mid-season stress is relieved at early pod filling. The capacity to produce a second flush of pods did not compensate fully for yield reduction due to water stress. Relief from mid-season stress also resulted in continued leaf production, N-2 fixation and vegetative biomass accumulation during pod filling. Despite the wide variation in the degree of change in vegetative biomass and N during pod filling, there were strong relationships between grain yield and net-above-ground biomass at maturity, and grain N and above-ground N at maturity. Only in the extreme situations were HI and nitrogen HI affected noticeably. In those treatments where there was a large second flush of pods, there was a pronounced biphasic pattern to pod number production, with HI also progressing through two distinct phases of increase separated by a plateau. The proportion of grain yield contributed to by biomass produced before pod filling varied from 0 to 61% with the contribution greatest under terminal water deficit. There was a larger effect of water deficit on N accumulation, and hence N-2 fixation, than on biomass accumulation. The study confirmed the applicability of a number of long-standing physiological concepts to the analysis of the effect of water deficit on mungbean, but also highlighted the difficulty of accounting for timing effects of water deficit where second flushes of pods alter canopy development, biomass and yield accumulation, and N dynamics. Crown Copyright (C) 2003 Published by Elsevier B.V. All rights reserved.

Growth performance and management of a mixed rainforest tree plantation

Monoculture plantations of Pinus, Eucalyptus and Acacia have been established oil rainforest lands throughout the world. However, this type of reforestation generally supplies low quality timber and contributes to landscape simplification. Alternatives to exotic monoculture plantations are now beginning to gain momentum with farmers and landholders attempting to establish a variety of rainforest trees in small plantations. When compared to the well studied commercial species, knowledge concerning the growth and management of many of these rainforest species is in its infancy. To help expand this limited knowledge base an experimental plantation of 16 rainforest tree species in a randomised design was established near Mt. Mee, in south-eastern Queensland, Australia. Changes in growth, form (based on stem straightness, branch size and branchiness), crown diameters and leaf area of each species were examined over 5 years. Patterns of height growth were also measured monthly for 31 months. Species in this trial could be separated into three groups based on their overall growth after 5 years and their growth patterns. Early successional status, low timber density, high maximum photosynthetic rates and large total leaf areas were generally correlated to rapid height growth. Several species (including Araucaria cunninghamii, Elaeocarpus grandis, Flindersia brayleyana, Grevillea robusta and Khaya nyasica) had above average form and growth, while all species in the trial had considerable potential to have increased productivity through tree selection. As canopy closure occurred at the site between years four and five, growth increments declined. To reduce stand competition a number of different thinning techniques could be employed. However, simple geometric or productivity based thinnings appear to be inappropriate management techniques for this mixed species stand as they would either remove many of the best performing trees or nearly half the species in the trial. Alternatively, a form based thinning would maintain the site's diversity, increase the average form of the plantation and provide some productivity benefits.

Seasonal patterns of growth, dehydrins and water-soluble carbohydrates in genotypes of Dactylis glomerata varying in summer dormancy

Background and Aims Summer dormancy in perennial grasses has been studied inadequately, despite its potential to enhance plant survival and persistence in Mediterranean areas. The aim of the present work was to characterize summer dormancy and dehydration tolerance in two cultivars of Dactylis glomerata (dormant 'Kasbah', non-dormant 'Oasis') and their hybrid using physiological indicators associated with these traits. Methods Dehydration tolerance was assessed in a glasshouse experiment, while seasonal metabolic changes which produce putative protectants for drought, such as carbohydrates and dehydrins that might be associated with summer dormancy, were analysed in the field. Key Results The genotypes differed in their ability to survive increasing soil water deficit: lethal soil water potential (ψ(s)) was -3(.)4 MPa for 'Kasbah' (although non-dormant), -1(.)3 MPa for 'Oasis', and -1(.)6 MPa for their hybrid. In contrast, lethal water content of apices was similar for all genotypes (approx. 0(.)45 g H2O g d. wt(-1)), and hence the greater survival of 'Kasbah' can be ascribed to better drought avoidance rather than dehydration tolerance. In autumn-sown plants, 'Kasbah' had greatest dormancy, the hybrid was intermediate and 'Oasis' had none. The more dormant the genotype, the lower the metabolic activity during summer, and the earlier the activity declined in spring. Decreased monosaccharide content was an early indicator of dormancy induction. Accumulation of dehydrins did not correlate with stress tolerance, but dehydrin content was a function of the water status of the tissues, irrespective of the soil moisture. A protein of approx. 55 kDa occurred in leaf bases of the most dormant cultivar even in winter. Conclusions Drought avoidance and summer dormancy are correlated but can be independently expressed. These traits are heritable, allowing selection in breeding programmes.

Growth responses of sugarcane to mycorrhizal spore density and phosphorus rate

Arbuscular mycorrhizal (AM) fungi, commonly found in long-term cane-growing fields in northern Queensland, are linked with both negative and positive growth responses by sugarcane ( Saccharum spp.), depending on P supply. A glasshouse trial was established to examine whether AM density might also have an important influence on these growth responses. Mycorrhizal spores ( Glomus clarum), isolated from a long-term cane block in northern Queensland, were introduced into a pasteurised low-P cane soil at 5 densities ( 0, 0.06, 0.25, 1, 4 spores/g soil) and with 4 P treatments ( 0, 8.2, 25, and 47 mg/kg). At 83 days after planting, sugarcane tops responded positively to P fertilizer, although responses attributable to spore density were rarely observed. In one case, addition of 4 spores/g led to a 53% yield response over those without AM at 8 mg P/kg, or a relative benefit of 17 mg P/kg. Root colonisation was reduced for plants with nil or 74 mg P/kg. For those without AM, P concentration in the topmost visible dewlap ( TVD) leaf increased significantly with fertiliser P (0.07 v. 0.15%). However, P concentration increased further with the presence of AM spores. Irrespective of AM, the critical P concentration in the TVD leaf was 0.18%. This study confirms earlier reports that sugarcane is poorly responsive to AM. Spore density, up to 4 spores/g soil, appears unable to influence this responsiveness, either positively or negatively. Attempts to gain P benefits by increasing AM density through rotation seem unlikely to lead to yield increases by sugarcane. Conversely, sugarcane grown in fields with high spore densities and high plant-available P, such as long-termcane-growing soils, is unlikely to suffer a yield reduction from mycorrhizal fungi.

Effects of high temperature on the growth and composition of sugarcane internodes

Sugarcane grown in the Ord River district of Western Australia has lower sucrose content than expected from earlier trials and experience in other irrigated districts. High temperatures have been hypothesised as a possible cause. The effects of high temperature (above 32 degrees C) on growth and carbon partitioning were investigated. A temperature regime of (25-38 degrees C) was compared with (23-33 degrees C). In one experiment, 7-month-old plants of cvv. Q117 and Q158 were subjected to the treatments for 2 months. In another experiment, the plants were allowed to regrow (ratoon) for 6 months. In both experiments, the higher temperature resulted in more, shorter internodes and higher moisture content. Most internodes from plants in the higher temperature treatment had lower sucrose content than internodes from the lower temperature. On a dry mass basis the internodes from the plants in the higher temperature had proportionately more fibre and hexoses but lower sucrose. Combined with an increased number of nodes in a stem of similar or shorter length this would result in higher stalk fibre and lower sucrose content. The data provided evidence that sugarcane partitions less carbon to stored sucrose when grown under high compared with low temperatures. The two cultivars partitioned carbon between soluble (sugars) and insoluble (fibre) fractions to different degrees. These experiments also indicate that the current models describing leaf appearance and perhaps sugarcane growth at temperatures above 32 degrees C, in general, need revision.

Nutrient deficiencies in greater yam and their effects on leaf nutrient concentrations

This paper describes the first systematic study of nutritional deficiencies of greater yam (Dioscorea alata). Yam plants (cv. 'Mahoa'a') were propagated from tuber discs and grown in nutrient solution, with nutrients supplied following a modified programmed nutrient-addition method. After an establishment period of four weeks, deficiencies of nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), sulfur (S), iron (Fe), boron (B), manganese (Mn), copper (Cu), zinc (Zn), and molybdenum (Mo) were induced by omitting the relevant nutrient from the solution. Foliar symptoms were recorded photographically. Notably, deficiencies of the mobile macronutrients failed to induce senescence of oldest leaves, while vine growth and younger leaves were affected. Leaf blades of the main stem were sampled in sequence and analyzed chemically, providing the distribution of each nutrient from youngest to oldest leaves in both adequately supplied and deficient plants. The nutrient-concentration profiles, together with the visible symptoms, indicated that little remobilization of mobile macronutrients had occurred. For both macro- and micronutrients, young leaves gave the best separation of nutrient concentrations between well-nourished and deficient plants.

Effect of physical, chemical and light treatments on germination and growth of tissue-cultured coconuts

The acclimatization and ex vitro establishment of tissue cultured coconut plantlets regenerated either from zygotic or somatic embryos could result to serious losses. Although high germination rates can be achieved in vitro, the survival of zygotic embryo derived plantlets in soil is very low (0-30%). Hence, treatments that could promote development of good quality seedlings having well-developed shoot and root is needed to increase seedling survival ex vitro. The effect of physical, chemical and light quality treatments on germination and growth of coconut embryos and tissue-cultured seedlings respectively, was investigated. The germination of coconut embryos was promoted when placed in a liquid Euwens (Y3) medium and incubated using a roller drum. Gibberellic acid (GA3) significantly affected growth of seedlings as it promoted shoot elongation, shoot and root expansion, and fresh and dry weight increase. However, GA3 did not significantly affect germination. In addition, the blue, red and yellow light significantly affected growth of seedlings as it promoted leaf and shoot elongation, fresh and dry weight increase, and root and leaf production. These conditions could be used to improve the growth and survival ex vitro of tissue cultured coconuts.

Towards developing reliable models of leaf area on grapevines (Vitis vinifera L.)

A field study in three vineyards in southern Queensland (Australia) was carried out to develop predictive models for individual leaf area and shoot leaf area of two cultivars (Cabernet Sauvignon and Shiraz) of grapevines (Vitis Vinifera L.). Digital image analysis was used to measure leaf vein length and leaf area. Stepwise regressions of untransformed and transformed models consisting of up to six predictor variables for leaf area and three predictor variables for shoot leaf area were carried out to obtain the most efficient models. High correlation coefficients were found for log10 transformed individual leaf and shoot leaf area models. The significance of predictor variables in the models varied across vineyards and cultivars, demonstrating the discontinuous and heterogeneous nature of vineyards. The application of this work in a grapevine modeling environment and in a dynamic vineyard management context are discussed. Sample sizes for quantification of individual leaf areas and areas of leaves on shoots are proposed based on target margins of errors of sampled data.