缺磷白羽扇豆排根与非排根区根尖分泌有机酸的比较

采用根系分泌有机酸原位收集方法及市郊液相色谱技术分析了供磷及缺磷后不同时间白羽扇豆（Ｌｕｐｉｎｕｓ ａｉｂｕｓ Ｌ．）非排根区根尖和排根分泌有机酸的种类和数量,以及相应的根尖、排根组织,茎木质部、韧皮部汁液中有机酸含量的变化。结果表明：⑴缺磷能够诱导白羽扇豆要系产生大量排根,根系的有机酸分泌量也明显增加。⑵无论在供磷或缺磷条件下,排根与非排根区根区根尖组织中的有机酸种类相同,但排根主要分泌柠檬酸和     

子叶磷在白羽扇豆缺磷适应性反应中的作用

实验用液体培养的方法,对比分析了在不同供磷条件下,白羽扇豆子叶中的磷对植物生长发育的影响,以及排根和根尖中有机酸积累和分泌的作用,结果表明,子叶中的磷能使白羽扇豆在完全缺磷２３d的环境中,不仅没有使干物质的积累减少,反而使干物质的积累略有增加,相反,如果没有子叶磷的供给,则使白羽扇豆在缺磷环境中产生强烈的抗胁迫反应,表现在干物质的积累明显下降,根系能产生大量的排根,排根能积累和分泌大量的柠檬酸,而根尖能积累和分泌萍果酸,在整个缺磷反应过程中,根尖中苹果酸的分泌要早于排根可柠檬酸的积累和分泌。     

6-BA对缺磷白羽扇豆排根形成和有机酸分泌的影响

缺磷条件下白羽扇豆能够形成排根,并增加有机酸分泌.但上述过程的调节机制尚不清楚.该文的结果表明,使用外源6-BA不影响缺磷白羽扇豆的生长和磷在体内的分配,但明显抑制了根簇的形成和有机酸分泌.经低浓度6-BA(10-8 mol/L)处理后转移至不含6-BA的缺磷营养液中继续培养的植株,其根簇形成和有机酸分泌得到恢复,甚至超过未经6-BA处理的缺磷植株;但高浓度6-BA(10-7 mol/L)对根簇形成和有机酸分泌的抑制作用不可恢复.对6-BA影响缺磷的白羽扇豆排根形成和有机酸分泌的可能机制进行了讨论.     

缺磷条件下白羽扇豆排根发育与生长素及miR164的关系

以缺磷条件下白羽扇豆为材料,观察了外源生长素NAA和生长素运输的抑制剂NPA 对白羽扇豆排根形成及其活性的影响,同时运用基因芯片与RT-PCR的方法分析了生长素信号转导途径中转录因子NAC1以及调控NAC1表达的上游microRNA164(miR164)在不同发育阶段排根中的表达变化,以探讨白羽扇豆在缺磷时排根形成与发育的调控机制.结果表明,缺磷胁迫下排根大量形成与生长素及其运输有关,排根NAC1的表达在初生阶段上调,成熟后下调,并受其上游的miR164的负调控,而排根衰老后则上述基因的表达都减弱.研究发现,在缺磷诱导的排根发生至发育成熟过程中,miR164、NAC1、生长素与排根发育之间很可能组成了一个级联系统,从而控制排根的发生与发育.     

缺磷或缺铁引起白羽扇豆锰中毒的机理

白羽扇豆在缺磷或缺铁条件下均有排根形式,并且根系还原力显著增加。缺磷、缺铁根系还原力在高峰期分别高于对照。缺磷与缺铁根系还原力高峰不仅出现的时期不同,而且还原力增加部位也不一样。缺磷处理的排根区具有很高的还原力,缺铁处理还原力较高的部位是在主根和侧根的根尖以及排根区。由于Ｍｎ＾４＋比Ｆｅ＾３＋更易被还原,致使根系还原力提高促使根际大量锰被还原,这是缺磷和缺铁造成白羽扇豆锰中毒的主要原因之一。     

缺磷或缺铁引起白羽扇豆锰中毒的机理

白羽扇豆在缺磷或缺铁条件下均有排根形成,并且根系还原力显著增加。缺磷、缺铁根系还原力在高峰期分别高于对照。缺磷与缺铁根系还原力高峰不仅出现的时期不同,而且还原力增加部位也不一样。缺磷处理的排根区具有很高的还原力,缺铁处理还原力较高的部位是在主根和侧根的根尖以及排根区。由于Ｍｎ４＋比Ｆｅ３＋更易被还原,致使根系还原力提高促使根际大量锰被还原,这是缺磷和缺铁造成白羽扇豆锰中毒的主要原因之一。     

磷和外源生长素对白羽扇豆(Lupinus albus L.)根形态和生理特性的影响

白羽扇豆(Lupinus albus L.)对磷和生长素表现出高度的根系形态和生理可塑性反应, 然而磷和生长素如何调节根形态和生理特性以及它们对根形态和生理的交互效应尚不清楚. 本研究通过水培实验旨在评价磷(0或250 μmol/L)和生长素(10^-8 mol/L NAA)对白羽扇豆根特性的影响及其交互效应. 结果表明, 缺少磷和生长素施用均明显改变了白羽扇豆的根形态(主根长度减少、一级侧根数目增加和排根大量形成)和生理特性(质子释放、柠檬酸分泌和酸性磷酸酶活性增加). 外源生长素的施用增加了缺磷白羽扇豆根系的响应度和敏感性. 磷和生长素对白羽扇豆根系形态和生理具有明显的交互作用. 主成分分析表明, 磷解释了根特性64.8%的信息, 而生长素解释了21.3%的信息, 表明磷供应对白羽扇豆根特性的影响比外源生长素施用更为重要. 白羽扇豆能够通过协调根系形态和生理对外部刺激(如缺磷和施用生长素)作出应答, 以及优化根形态和生理之间的关系从而最大化获取磷素资源.     

不同磷效率小麦品种对缺磷胁迫反应的比较

在营养液培养条件下,以根据相对产量为指标筛选出的6个不同磷效率的小麦(Triticum aestivum L.)品种为材料,对其苗期在缺磷条件下生长、根冠磷含量及其分配,以及叶片韧皮部汁液中磷浓度等进行了比较研究。结果表明,缺磷抑制植株地上部生长,但刺激根系生长,导致植株根／冠比增加。无论在供磷或缺磷条件下,磷高效品种的根冠生长速率都低于磷低效品种。缺磷导致植株体内的磷含量下降与根系相比,地上部磷含量的下降速率更快。但在缺磷条件下,不同磷效率的小麦品种根冠间的磷分配变化没有差异。研究发现,在正常供磷条件下,磷高效小麦品种的叶片韧皮部汁液中磷浓度较低,而磷低效品种的叶片韧皮部汁液中磷浓度较高。但开始缺磷后,磷高效品种的叶片韧皮部汁液中的磷浓度下降较慢,使其相对磷浓度较高。缺磷后10天,磷低效品种叶片韧皮部汁液中的磷浓度为供磷对照的35．9％,而磷高效品种叶片韧皮部汁液中的磷浓度为供磷对照的59％。     

排根的形成及其所分泌的有机酸的调节

排根是在磷、铁、氮等养分缺乏条件下形成的一种特殊根系结构,能够形成排根的植物种类有限.不同植物排根分泌的有机酸种类和数量可能不同,如在缺磷条件下白羽扇豆的排根所释放的柠檬酸可达植物总干重的23%,使根簇周围的柠檬酸浓度达50～90 μmol*g-1土壤.这是一种防止胞质过度酸化以及柠檬酸过度积累而超过液泡储存能力的解毒机制.其分泌可能受阴离子通道的调控.     

Root Cluster Formation and Citrate Exudation of White Lupin （Lupinus albus L.） as Related to Phosphorus Availability

A split-root system was used to investigate whether the external or internal P concentration controls root cluster formation and citrate exudation in white lupin (Lupinus albus L.) grown under controlled conditions. In spite of low P concentrations in the shoots and roots of the -P plant, its dry weight was not reduced compared with the P plant. Supplying external P (0.25 mmol/L) to one root half resulted in an increase in P concentration not only in the shoot, but also in the P-deprived root half, indicating P cycling within the plants. Omitting P from both split-root pots stimulated root cluster formation in both root halves,whereas P supply to one root half stimulated root cluster formation at the beginning of the treatment. Neither P supply to just one root half continuously nor resupply of P to one root half after 19 d of P starvation inhibited root cluster formation on the P-deprived side, although the concentration of P in this root half and shoot increased markedly. The results indicate that root cluster formation in L. albus is controlled by both shoot and root P concentrations. The rates of citrate exudation by both root halves with P deficiency were higher than those of the one root half supplied with P only. In the treatment with one root half supplied with P, the rates of citrate exudation by either the P-supplied or -deprived root halves were almost the same,regardless of P concentration in the roots. The results suggest that internal P concentration controls root cluster formation and citrate exudation in white lupin, but these processes may be regulated by different mechanisms.     

Root Cluster Formation and Citrate Exudation of White Lupin (Lupinus albus L.) as Related to Phosphorus Availability

A split-root system was used to investigate whether the external or internal P concentration controls root cluster formation and citrate exudation in white lupin (Lupinus albus L.) grown under controlled conditions. In spite of low P concentrations in the shoots and roots of the -P plant, its dry weight was not reduced compared with the P plant. Supplying external P (0.25 mmol/L) to one root halfresulted in an increase in P concentration not only in the shoot, but also in the P-deprived root half, indicating P cycling within the plants. Omitting P from both split-root pots stimulated root cluster formation in both root halves,whereas P supply to one root halfstimulated root cluster formation at the beginning of the treatment. Neither P supply to just one root half continuously nor resupply of P to one root half after 19 d of P starvation inhibited root cluster formation on the P-deprived side, although the concentration of P in this root half and shoot increased markedly. The results indicate that root cluster formation in L. albus is controlled by both shoot and root P concentrations. The rates of citrate exudation by both root halves with P deficiency were higher than those of the one root half supplied with P only. In the treatment with one root half supplied with P, the rates of citrate exudation by either the P-supplied or -deprived root halves were almost the same,regardless of P concentration in the roots. The results suggest that internal P concentration controls root cluster formation and citrate exudation in white lupin, but these processes may be regulated by different mechanisms.     

Distribution and Function of Proteoid Roots and other Root Clusters

Proteoid roots are bottlebrush-like clusters of rootlets which form along lateral roots. They are characteristic of most species of the Proteaceae, which are mainly distributed in Australia and South Africa. Homologous root clusters are present in species of the Casuarinaceae, Mimosaceae, Fabaceae, Myricaceae and Moraceae. Many similarities exist between these species in relation to morphology and function of root clusters. Many are non-mycorrhizal and are highly efficient in phosphorus (P) acquisition. In these species, proteoid roots and proteoid-like root clusters are abundant when grown on infertile soils. Their formation is predominantly affected by the P status of the plants, being induced at low P levels and repressed at high P levels. Proteoid roots and proteoid-like root clusters play an important role in acquisition of P and other mineral nutrients. Although increase in root surface area may be a contributing factor, in many species these roots excrete large amounts of organic acids and phenolics. The excretion of these compounds in a small soil volume gives rise to extensive nutrient mobilization by acidification, reduction and chelation of sparingly soluble forms of P and micronutrients such as Fe and Mn.     

Acid phosphatase activity in phosphorus-deficient white lupin roots

White lupin ( Lupinus albus L.) develops proteoid roots when grown in phosphorus (P)-deficient conditions. These short, lateral, densely clustered roots are adapted to increase P availability. Previous studies from our laboratory have shown proteoid roots have higher rates of non-photosynthetic carbon fixation than normal roots and altered metabolism to support organic acid exudation, which serves to solubilize P in the rhizosphere. The present work indicates that proteoid roots possess additional adaptations for increasing P availability and possibly for conserving P in the plant. Roots from P-deficient (–P) plants had significantly greater acid phosphatase activity in both root extracts and root exudates than comparable samples from P-sufficient (+P) plants beginning 10 d after emergence. The increase in activity in –P plants was most pronounced in the proteoid regions. In contrast, no induction of phytase activity was found in –P plants compared to +P plants. The number of proteoid roots present was not affected by the source of phosphorus supplied, whether organic or inorganic forms. Adding molybdate to the roots increased the number of proteoid roots in plants supplied with organic P, but not inorganic P. Increased acid phosphatase activity was detected in root exudates in the presence of organic P sources. Native-polyacrylamide gel electrophoresis demonstrated that under P-deficient conditions, a unique isoform of acid phosphatase was induced between 10 and 12 d after emergence. This isoform was found not only within the root, but it comprised the major form exuded from proteoid roots of –P plants. The fact that exudation of proteoid-root-specific acid phosphatase coincides with proteoid root development and increased exudation of organic acids indicates that white lupin has several coordinated adaptive strategies to P-deficient conditions.     

Effect of phosphorus supply on the formation and function of proteoid roots of white lupin (Lupinus albus L.)

We investigated (1) the effect of constant and altered inorganic phosphate (P_i) supply (1–100 mmol m^–3) on proteoid root production by white lupin ( Lupinus albus L.); and (2) the variation in citrate efflux, enzyme activity and phosphate uptake along the proteoid root axis in solution culture. Proteoid root formation was greatest at P_i solution concentrations of 1–10 mmol m^–3 and was suppressed at 25 mmol m^–3 P_i and higher. Except at 1 mmol m^–3 P_i, the formation of proteoid roots did not affect plant dry matter yields or shoot to root dry matter ratios, indicating that proteoid roots can form under conditions of adequate P supply and not at the expense of dry matter production. Plants with over 50% of the root system as proteoid roots had tissue P concentrations considered adequate for maximum growth, providing additional evidence that proteoid roots can form on P-sufficient plants. There was an inverse relationship between the P_i concentration in the youngest mature leaf and proteoid root formation. Citrate efflux and the activities of enzymes associated with citric acid synthesis (phosphoenolpyruvate carboxylase and malate dehydrogenase) varied along the proteoid root axis, being greatest in young proteoid rootlets of the 1–3 cm region from the root tip. Citrate release from the 0–1 and 5–9 cm regions of the proteoid root was only 7% (per unit root length) of that from the 1–3 cm segment. Electrical potential and ³²P_i uptake measurements showed that P_i uptake was more uniform along the proteoid root than citrate efflux.     

Isoflavonoid exudation from white lupin roots is influenced by phosphate supply, root type and cluster-root stage

The internal concentration of isoflavonoids in white lupin (Lupinus albus) cluster roots and the exudation of isoflavonoids by these roots were investigated with respect to the effects of phosphorus (P) supply, root type and cluster-root developmental stage.To identify and quantify the major isoflavonoids exuded by white lupin roots, we used high-pressure liquid chromatography (HPLC) coupled to electrospray ionization (ESI) in mass spectrometry (MS).The major exuded isoflavonoids were identified as genistein and hydroxygenistein and their corresponding mono- and diglucoside conjugates. Exudation of isoflavonoids during the incubation period used was higher in P-deficient than in P-sufficient plants and higher in cluster roots than in noncluster roots. The peak of exudation occurred in juvenile and immature cluster roots, while exudation decreased in mature cluster roots.Cluster-root exudation activity was characterized by a burst of isoflavonoids at the stage preceding the peak of organic acid exudation. The potential involvement of ATP-citrate lyase in controlling citrate and isoflavonoid exudation is discussed, as well as the possible impact of phenolics in repelling rhizosphere microbial citrate consumers.