Few studies have examined if perennial leguminous cover crops are able to transfer nitrogen (N) via common mycorrhizal networks (CMNs) to neighboring fruit trees; the gradient of such N transfer ...could affect the N nutrition of both plants. Using separated three-column chambers to grow plants in a greenhouse, 99 atom% 15N as (15NH4)2SO4 was applied to leaves of white clover (Trifolium repens L.) and 15N was then traced in neighboring citrus (Citrus sinensis (L.) Osbeck) seedlings interconnected by an arbuscular mycorrhizal fungus (AMF, Rhizophagus intraradices). A range of 66.85–68.74% mycorrhizal colonization in white clover (mycorrhizal and/or Rhizobium trifolii inoculated) and 19.29–23.41% in citrus (non-mycorrhizal inoculated) was observed after 12 months of AMF inoculation in the white clover, indicating a successful CMN linkage was established between these two plant species. This CMN establishment resulted in significant increases in biomass, N accumulation, and 15N content of citrus when accompanied with nodulated and mycorrhizal fungus colonized white clover. N transfer from white clover to citrus was significantly greater under nodulation plus mycorrhization (46.23 mg N per pot, 1.71% of N transferred) than under non-inoculated control (4.36 mg N per pot, 0.21% of N transferred), and higher than sole mycorrhization (36.34 mg N per pot, 1.42% of N transferred). The percentage of N in citrus derived from white clover under nodulated/mycorrhization was 1.83–1.93%, and was highest in leaves (3.31%), moderate in stems (2.47%), and lowest in roots (0.41%) of citrus. In summary, results from this experiment demonstrated that nearly 2.0% of N transferred from white clover to citrus via CMN. Further studies are needed to quantify N transfer between white clover and citrus by other routes, including soil or root exudation pathways.
Spermidine (Spd) may be involved in the regulation of plant adaptation to drought stress. The objectives of the study were to identify the physiological effect and elucidate the possible mechanism ...caused by exogenous Spd (0.05 mM) in white clover under water stress induced by 20% polyethylene glycol 6000 for 12 days. Water stress elevated significantly the accumulation of reactive oxygen species and malonaldehyde, and resulted in the decrease of cell membrane stability, relative water content and relative growth mte. Spd effectively alleviated the damage effect from water stress. Spd-treated plants showed a promoted the ascorbate-glutathione cycle and maintained greater antioxidant enzyme activities (superoxide dismutase, peroxidase and catalase), as well as higher transcript level of genes encoding antioxidant enzymes than untreated plants. Additionally, the plants treated with Spd under water stress exhibited more accumulated organic solutes including soluble sugar, reducing sugar, betaine and free proline. Spd also accelemted proline catabolism and biosynthesis proceeding from glutamate pathway during water stress, but had no effect on the omithine pathway of proline biosynthesis. These results suggest that exogenous Spd under water stress may directly or indirectly regulate antioxidant defense system, organic solutes accumulation and proline metabolism.
The objective of this present investigation was to study the different response on drought tolerance and post-drought recovery for two different genotypes of white clover (a drought-tolerant ...small-leafed white clover and a drought-sensitive large-leafed white clover) associating with antioxidative enzyme and lignin metabolism. Two materials were exposed to 12 days of drought stress and 6 days of rewatering in a greenhouse. Significantly higher activities of superoxide dismutase, ascorbate peroxidase, catalase, guaiacol peroxidase and root activity as well as lower electrolyte leakage and malondialdehyde levels of the small-leafed in leaves and roots were observed as compared to the large-leafed under drought stress and rewatering. Lignification-related enzymes (polyphenol oxidase, cinnamyl alcohol dehydrogenase) activities of two materials in leaves gradually increased from 0 to 9 days of drought stress; then declined at 12 days of drought stress and after rewatering. After 12 days of drought stress, lignin content of two materials in roots was significantly higher relative to the well-watered. Lignin content of leaves in the small-leafed also increased significantly after 6 days of rewatering, but not in the large-leafed. The small-leafed exhibited more sensitive lignin metabolism and faster lignification in leaves and roots than the large-leafed regarding the response of drought stress. These results indicated that antioxidant efficiency and lignin metabolism played important roles in response to drought stress and rewatering for different white clovers. Comparatively higher reactive oxygen species scavenging system and more sensitive lignin metabolism could be associated with better drought tolerance and post-drought recovery in white clover.
For pastures, root turnover can have an important influence on nutrient and carbon cycling, and plant performance. Turnover was calculated from mini‐rhizotron observations for chicory (Cichorium ...intybus), lucerne (Medicago sativa), perennial ryegrass (Lolium perenne) and white clover (Trifolium repens) grown in the Manawatu, New Zealand. The species were combined factorially with four earthworm species treatments and a no‐earthworm control. Split plots compared the effects of not cutting and cutting the shoots at intervals. Observations were made c. 18 days apart for 2.5 years. This article concentrates on differences between plant species in root turnover in the whole soil profile to 40 cm depth. At this scale, earthworm effects were generally small and short lived. For ryegrass and white clover, root length and mass were linearly related (R² = 0.82–0.99). For chicory and lucerne, the relationships were poorer (R² = 0.38–0.77), so for those species length turnover may be a poor indicator of mass turnover. Standing root length, total growth and death generally decreased in the sequence ryegrass > lucerne > chicory = white clover. In length terms, scaled turnover (growth divided by average standing root length) generally followed the sequence lucerne > white clover > perennial ryegrass = chicory. Across species the scaled turnover rate averaged 3.4 per year or 0.9% per day. Cutting shoots reduced standing root length, growth and death, but increased scaled turnover. These results indicate fast and prolonged root turnover. For ryegrass and white clover, at least there is need to reappraise how to measure and model shoot : root ratios, dry matter production and carbon cycling.
The effect of calcium (Ca²⁺) on Trifolium repens L. seedlings subjected to cadmium (Cd²⁺) stress was studied by investigating plant growth and changes in activity of antioxidative enzymes. ...Physiological analysis was carried out on seedlings cultured for 2 weeks on half-strength Hoagland medium with Cd²⁺ concentrations of 0, 400 and 600 μM, and on corresponding medium supplied with CaCl₂ (5 mM). Exposure to increasing Cd²⁺ reduced the fresh weight of the upper part (stems + leaves) of the seedlings more strongly than that of the root system. In both parts of T. repens seedlings H₂O₂ level and lipid peroxidation increased. In the upper part, Cd²⁺ exposure led to a significant decrease in the activity of superoxide dismutase, catalase and glutathione peroxidase and an increase in ascorbate peroxidase activity. In contrast, the roots showed an increase in the activity of antioxidative enzymes under Cd²⁺ stress. Ca²⁺ addition to medium reduced the Cd²⁺ accumulation, and considerably reversed the Cd²⁺-induced decrease in fresh mass as well as the changes in lipid peroxidation in the both parts of T. repens seedlings. Ca²⁺ application diminished the Cd²⁺ effect on the activity of antioxidative enzymes in the upper part, even though it did not significantly affect these enzymes in the roots. So the possible mechanisms for the action of Ca²⁺ in Cd²⁺ stress were considered to reduce Cd²⁺ accumulation, alleviate lipid peroxidation and promote activity of antioxidative enzymes.
Various nitric oxide (NO) regulators including the NO donor sodium nitroprusside (SNP), the NO scavenger 2-(4-carboxyphenyl)-4,4,5,5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO), the NO-synthase ...inhibitor
N
G
-nitro-
L
-Arg-methyl ester (
L
-NAME), and the SNP analogues sodium nitrite/nitrate and sodium ferrocyanide were investigated to elucidate the role of NO in white clover (
Trifolium repens
L.) plants after long-term (5 days) exposure to cadmium (Cd). A dose of 100 μM Cd stress significantly restrained plant growth and decreased the concentrations of chlorophyll and NO in vivo, whereas it disrupted the balance of stress-related hormones and enhanced the accumulation of Cd, thereby inducing reactive oxygen species (ROS) burst. However, the inhibition of plant growth was relieved by 50 μM SNP through its stimulation of ROS-scavenging compounds (ascorbic acid, ascorbate peroxidase, catalase, glutathione reductase, non-protein thiol, superoxide dismutase, and total glutathione), regulation of H
+
-ATPase activity of proton pumps, and increasing jasmonic acid and proline but decreasing ethylene in plant tissues. Even so, the alleviating effect of SNP on plant growth was counteracted by cPTIO and
L
-NAME and was not observed with SNP analogues, suggesting that the protective roles of SNP are related to the induction of NO. These results suggest that NO may improve the Cd tolerance of white clover plants by eliminating oxidative damage, re-establishing ATPase activity, and maintaining hormone equilibrium. Improving our understanding of the role of NO in white clover plants is key to expanding the plantations to various regions and the recovery of pasture species in the future.
Optimum plant densities are a key to maximise yields in most crops. However, such information is often lacking for more environmentally sound cropping systems, such as living mulches (LM) for small ...grains. In 2004 and 2005, three trials were conducted in the Swiss Midlands on fields managed in accordance with the Swiss organic farming guidelines. The objective of the study was to determine whether seeding density of winter wheat (
Triticum aestivum L.) is a relevant factor for determining grain yield in a white clover (
Trifolium repens L.) living mulch. The winter wheat cv. Titlis was directly sown in wide spaced rows (0.375
m) at densities of 300 (LM300), 450 (LM450) or 600 (LM600) viable grains m
−2 in a white clover living mulch established at a seeding rate of 15
kg
ha
−1. A bare soil control treatment with a wheat density of 450 viable grains m
−2 (BS450) was also included in the trials. Mean grain yields of LM300, LM450, and LM600 never reached the values observed in BS450. This was mainly due to a lower ear density, which, nevertheless, increased linearly with the seeding density within the living mulch in all trials, but the rate of increase depended on the environment. The decrease of the grain weight brought about by the increasing seeding density had only a marginal impact on the grain yield, which was increased from 1.31, 1.98, and 4.09
Mg
ha
−1 (LM300) to 1.97, 2.64, and 4.75
Mg
ha
−1 (LM600) for each of the three trials in the study. Significantly higher protein contents were observed for LM300 compared to the higher densities in the living mulch and to BS450. Our research showed that an increase of the seeding density is an effective mean to increase the grain yield in living mulch systems with white clover. However, it is likely that the control of the living mulch to reduce competition with the main crop is a more relevant factor.
O trevo branco é utilizado no Sul do Brasil conferindo qualidade nas pastagens ao qual se encontra, podendo ser maximizada a sua produção pelo manejo de macro e micronutrientes. Porém, alguns ...micronutrientes como o boro, possuem limites estreitos entre a deficiência e a toxicidade. Dessa forma, esse trabalho teve por objetivo avaliar os efeitos da aplicação de doses de boro sobre a produção de forragem e componentes radiculares de trevo branco cultivadas em um Latossolo Vermelho distroférrico. O experimento foi conduzido em casa de vegetação da Universidade Tecnológica Federal do Paraná, localizada no Município de Pato Branco - PR. O delineamento experimental foi inteiramente casualizado, com cinco repetições e os tratamentos consistiram de cinco níveis de boro (0; 0,5; 1; 2 e 4 kg ha-1). Foram avaliadas as características referentes à produção de raízes, nodulação e matéria seca da parte aérea em plantas de trevo branco. A adubação aumentou a produção de matéria seca de raízes, o número de nódulos e a matéria seca de nódulos até 2,8 kg de boro ha-1, resultando em maior produção de matéria seca de forragem. As maiores taxas, no entanto, apresentaram efeitos negativos sobre a produção de trevo branco, devido à sua toxicidade.
Four breeding lines (BLs) of white clover, which range in terms of their disposition to increase biomass in response to low or high phosphate (P) levels in soil, were evaluated further when grown ...hydroponically. BL 45 increased both fresh and dry weight in response to P-deprivation, while BL 49 displayed the same significant stimulation in response to sufficient P. However, when compared these lines did not accumulate any significantly higher levels of P, and all four lines showed the same changes to root:shoot ratio in response to P-deprivation, as well as the induction of both soluble and cell-wall-associated acid phosphatase activity. These results confirm that there is no direct relationship between growth (as an increase in biomass) and acid phosphatase activity and further some responses to P-deprivation may be common to all genetic backgrounds suggesting that they are not part of the intricate mechanisms governing these responses.
Antioxidant response to drought in red (Trifolium pratense L., cv. “Start”) and white clover (Trifolium repens L, cv. “Haifa” and cv. “Debut”) grown as soil cultures was evaluated in water-deprived ...and recovered plants. Drought provoked oxidative stress in leaves confirmed by the considerable changes in electrolyte leakage, malondialdehyde, hydrogen peroxides and proline contents. Immunoblot of Δ-1-pyrroline-5-carboxylate synthetase (P5CS), which catalyzes the first two steps in proline biosynthesis, revealed strong induction of the enzyme in red clover plants submitted to drought. Water-deprived white clover plants exhibited distinct P5CS profiles. This was related to different drought tolerance of the studied T. repens cultivars. Isoenzyme analyses of superoxide dismutase (SOD), peroxidase (POX) and catalase (CAT) demonstrated certain differences in antioxidant defence among the tested varieties. It was confirmed that MnSOD (in both T. repens and T pratense) and FeSOD (in T. repens) isoforms were the most affected by drought. The red clover cultivar “Start” exhibited the lowest FeSOD and POX activities which could contribute to its poor performance under water deprivation.
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