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1.
白梭梭同化枝对干旱胁迫的生理生态响应   总被引:8,自引:1,他引:7  
对白梭梭同化枝自然水分胁迫下的季节抗旱特征进行研究。结果表明:自然干旱胁迫条件下,不同生长发育时期同化枝的主导抗旱生理因子不同。5月31日至6月29日盛花期间,同化枝内源ABA含量甚微,气孔导度值与光合强度处于生长发育过程的最高水平,可溶性糖含量呈现下降趋势,而脯氨酸含量略有增加,植物通过渗透调节作用适应此阶段轻度干旱胁迫;6月29日至7月26日盛夏期间,ABA迅速积累,气孔导度值降为生长发育过程的最低值,叶绿素分解,可溶性糖与脯氨酸均呈现快速积累趋势,且可溶性糖积累强度大于脯氨酸;8月9日至8月22日同化枝生长发育后期,ABA急剧积累为生长发育过程的最高浓度,气孔导度值有所增大,脯氨酸和可溶性糖保持在高水平平稳变化。此阶段高浓度ABA调节植物生理过程适应干旱的效应受CTK、IAA两种内源激素的抑制,进而抑制脯氨酸和可溶性糖的继续积累。  相似文献   

2.
We investigated effects of water stress and external abscisic acid (ABA) supply on shoot growth, stomatal conductance and water status in 1-year-old cuttings of a drought-sensitive poplar genotype Populus x euramericana cv. I-214 (Italica) and a drought-tolerant genotype P. 'popularis 35-44' (popularis). Populus popularis was more productive and maintained higher leaf water potentials throughout the drought treatment than cv. Italica. Supply of ABA to the xylem sap caused a greater decline in growth and more leaf abscission in shoots of cv. Italica than in shoots of P. popularis. Immediately after initiation of the drought treatment in P. popularis, the ABA concentration ([ABA]) of the xylem increased rapidly and stomatal conductance declined; however, stomatal conductance had returned to control values by the third day of the drought treatment, coincident with a gradual decline in xylem [ABA]. In contrast, xylem [ABA] of cv. Italica initially increased more slowly than that of P. popularis in response to the drought treatment, but the increase continued for 3 days at which time a tenfold increase in xylem [ABA] was observed that was followed by abscission of more than 40% of the leaves. We conclude that sensitivity of poplar roots to variation in soil water content varies by clone and that a rapid short-term accumulation of ABA in shoots in response to water stress may contribute to drought tolerance.  相似文献   

3.
Tree peony is well known and sought after for its large, colorful flowers. Its propagation is via vegetative methods. Mech-nisms of the adventitious rooting and the regulation of rooting processes are the principles and techniques of plant propagation and improvement. Microstructures and fluctuations of phytohormones in the adventitious rooting were studied with the etiolated softwood shoots of Paeonia suffruticosa 'Yinfen Jinlin'. There are no pre-primordia in the shoots of the cultivar. Adventitious roots are produced in five stages: shoot selection, primordium initiation, primordium growth, conducting tissue differentiation and root protrusion. Primordia initiated in the cortex. The contents of the endogenous hormones, IAA, ABA and GA, were 5.842, 0.873 and 1.043 nmol·g-1 FW on the bases of shoots, respectively. CTKs which included isopentenyl adenine (iPA), zeatin riboside (ZR) and dihy-drozeatin riboside (DHZR) were 0.949, 0.695 and 2.034 nmol·g-1 FW, respectively. DHZR is active among CTKs. The ratio of IAA to GA, CTK and ABA clearly increased at the stage of primordium initiation, while they showed low levels at the stages of primordium growth. The ratios were restored at the shoot levels at the stage of root protrusion. IBA provoked primordia initiation in the cortex, the vascular cambium, the pith and even in the callus induced on the base of shoots. IAA levels in the treated shoots increased gradually to its highest level (three times of control) at the stage of conducting tissue differentiation. The ratios of IAA to GA, CTK and ABA clearly decreased at the stage of primordium initiation. The ratio of IAA to ABA is regulated at 10:1.  相似文献   

4.
为开发朝仓山椒的椒芽菜价值,采用酶联免疫等方法,在休眠至萌发的过程,测定了枝条的ABA、IAA、GA3、ZR的含量以及可溶性蛋白、可溶性糖、游离氨基酸和游离脯氨酸的动态变化。结果表明,4种内源激素在芽内的含量存在极明显的差异,排序为ABA〉IAA〉GA3〉ZR;测定期内,IAM/ABA呈现较明显的上升趋势,GA3/ABA表现为较平缓的“升-降-升”趋势,IAA+GA3+ZR/ABA呈现出剧烈的“升-降-升”趋势;可溶性蛋白含量表现为“先升后降”,可溶性糖、游离氨基酸和游离脯氨酸含量表现为稳定的下降趋势。  相似文献   

5.
Tree peony is well known and sought after for its large, colorful flowers. Its propagation is via vegetative methods. Mechnisms of the adventitious rooting and the regulation of rooting processes are the principles and techniques of plant propagation and improvement. Microstructures and fluctuations of phytohormones in the adventitious rooting were studied with the etiolated softwood shoots of Paeonia suffruticosa ‘Yinfen Jinlin’. There are no pre-primordia in the shoots of the cultivar. Adventitious roots are produced in five stages: shoot selection, primordium initiation, primordium growth, conducting tissue differentiation and root protrusion. Primordia initiated in the cortex. The contents of the endogenous hormones, IAA, ABA and GA, were 5.842, 0.873 and 1.043 nmol•g–1 FW on the bases of shoots, respectively. CTKs which included isopentenyl adenine (iPA), zeatin riboside (ZR) and dihydrozeatin riboside (DHZR) were 0.949, 0.695 and 2.034 nmol•g–1 FW, respectively. DHZR is active among CTKs. The ratio of IAA to GA, CTK and ABA clearly increased at the stage of primordium initiation, while they showed low levels at the stages of primordium growth. The ratios were restored at the shoot levels at the stage of root protrusion. IBA provoked primordia initiation in the cortex, the vascular cambium, the pith and even in the callus induced on the base of shoots. IAA levels in the treated shoots increased gradually to its highest level (three times of control) at the stage of conducting tissue differentiation. The ratios of IAA to GA, CTK and ABA clearly decreased at the stage of primordium initiation. The ratio of IAA to ABA is regulated at 10:1.  相似文献   

6.
通过对不同产量类型雷竹林植株及竹笋激素的分析,认为GA3和ABA调控着雷竹鞭侧芽的萌发,也就是影响着竹林的产量,而IAA和CTK对雷竹鞭侧芽的萌发没有明显的调控作用,但影响着竹笋的生长发育;通过对氨基酸含量的分析认为高、中、低雷竹林竹笋的氨基酸含量基本相同,但覆盖雷竹林的春季竹笋氨基酸含量明显下降;通过对竹笋营养成分的分析,得出蛋白质、N、P、K含量从高、中、低产林呈下降趋势,并还认为竹林中适当增施N肥有利于笋产量的提高。  相似文献   

7.
为研究MS培养基不同浓度的KNO_3对猕猴桃离体繁育的影响,以‘红阳’猕猴桃组培幼苗为试验材料,就培养基中添加不同浓度的KNO_3改良MS培养基对猕猴桃增殖的新梢形态,抗氧化酶活性、苯丙氨酸解氨酶活性及内源激素含量变化的影响进行了试验研究。结果表明:当KNO_3质量浓度从1 140 mg/L增加至2 660 mg/L时,猕猴桃幼苗中的抗氧化酶(SOD、POD和CAT)活性、PAL活性、内源激素(IAA、GA_3和CTK)含量、IAA/ABA、GA_3/ABA和CTK/ABA均呈先增大后减小的变化趋势;ABA含量则随KNO_3质量浓度的增大呈先减小后增大的趋势。当KNO_3质量浓度为1 900 mg/L时,体内氧化酶、苯丙氨酸解氨酶活性最高,内源激素IAA、GA_3、CTK含量较高,ABA含量较低,植株发育正常。  相似文献   

8.
Perks MP  Irvine J  Grace J 《Tree physiology》2002,22(12):877-883
We investigated the effect of drought on canopy stomatal conductance (g(c)), and examined the hypothesis that g(c) is controlled by the chemical messenger abscisic acid (ABA) produced in roots. Beginning in November 1994, we subjected a mature stand of Scots pine (Pinus sylvestris L.) to an imposed 11-month drought. Control plots were maintained at average-season soil water content. Xylem sap was extracted from shoots at regular intervals from April to November 1995. Soil water, sap flow and leaf water potentials (predawn to dusk) were recorded at the same time. Canopy stomatal conductance was calculated from sap flow data and xylem sap ABA concentrations ([ABA(xyl)]) were measured by radioimmunoassay. Mean [ABA(xyl)] in control trees was 250 micromol m(-3). No diurnal variation in [ABA(xyl)] was detected. With soil drying, [ABA(xyl)] increased to a maximum in summer (600 micromol m(-3)), but decreased again toward autumn; however, no significant increase in ABA flux to the leaves occurred. A decline in g(c) was detected when volumetric soil water content declined below 0.12. The decline in g(c) could not have been mediated by increasing [ABA(xyl)] because stomatal closure appeared to precede any increase in [ABA(xyl)]. Peak sap flow velocity data were used to estimate delivery times for root-to-shoot signals in 15-m tall trees. Under normal field conditions, a signal would take 12 days to travel from the site of production (roots) to the presumed site of action (shoots). However, under drought conditions it may take a chemical signal in excess of 6 weeks. We conclude that a feedforward model of short-term stomatal response to soil drying, based solely on the action of a chemical messenger from the roots, is not applicable in mature conifer trees because signal transmission is too slow.  相似文献   

9.
以当年生盆栽美国红枫扦插苗为试验材料,采用不同浓度的脱落酸(ABA)溶液喷施美国红枫扦插苗,测定干旱胁迫下植株的相对电导率、脯氨酸(Pro)、丙二醛(MDA)及可溶性糖含量等各项生理指标,研究外源喷施ABA对美国红枫抗旱性的影响。结果表明,在干旱条件下,外源喷施ABA可以缓解其伤害程度,促进美国红枫叶片中脯氨酸和可溶性糖含量的增加,抑制MDA的积累,延缓叶片相对电导率上升,其中以10 mg/L浓度ABA处理效果最明显。因此,干旱胁迫下,喷施一定浓度的脱落酸可有效提高美国红枫的抗旱能力,维持其正常生理代谢。  相似文献   

10.
Pinus radiata D. Don is one of the most abundant species in the north of Spain. Knowledge of drought response mechanisms is essential to guarantee plantation survival under reduced water supply as predicted in the future. Tolerance mechanisms are being studied in breeding programs, because information on such mechanisms can be used for genotype selection. In this paper, we analyze the changes of leaf water potential, hydraulic conductance (K(leaf)), stomatal conductance and phytohormones under drought in P. radiata breeds (O1, O2, O3, O4, O5 and O6) from different climatology areas, hypothesizing that they could show variable drought tolerance. As a primary signal, drought decreased cytokinin (zeatin and zeatin riboside-Z?+?ZR) levels in needles parallel to K(leaf) and gas exchange. When Z?+?ZR decreased by 65%, indole-3-acetic acid (IAA) and abscisic acid (ABA) accumulation started as a second signal and increments were higher for IAA than for ABA. When plants decreased by 80%, Z?+?ZR and K(leaf) doubled their ABA and IAA levels, the photosystem II yield decreased and the electrolyte leakage increased. At the end of the drought period, less tolerant breeds increased IAA over 10-fold compared with controls. External damage also induced jasmonic acid accumulation in all breeds except in O5 (P. radiata var. radiata?×?var. cedrosensis), which accumulated salicylic acid as a defense mechanism. After rewatering, only the most tolerant plants recovered their K(leaf,) perhaps due to an IAA decrease and 1-aminocyclopropane-1-carboxylic acid maintenance. From all phytohormones, IAA was the most representative 'water deficit signal' in P. radiata.  相似文献   

11.
We conducted field and pot experiments to investigate the effects of brassinolide on 1-year-old Robinia pseudoacacia L. seedlings. In the field experiment, seedling roots were soaked in brassinolide solutions containing 0–0.4 mg/l pure brassinolide before planting. Survival and growth of the seedlings were determined 8 months later. The results showed that soaking roots in brassinolide prior to planting significantly increased the survival and growth of seedlings. The best results were in the 0.2 mg/l brassinolide treatment. In the pot experiment, roots were soaked in 0–0.4 mg/l brassinolide before planting followed by a foliar application of brassinolide when the seedlings leafed out. After the seedlings were established, the soil water content in the pots was regulated to simulate drought conditions and various physiological parameters were measured. The results showed that treatment with 0.2 mg/l brassinolide decreased the transpiration rate, stomatal conductance and malondialdehyde (MDA) content of seedlings growing under moderate or severe water stress compared to untreated seedlings. Leaf water content, predawn water potential, soluble sugar content, free proline content, and superoxide dismutase (SOD), peroxidase (POD) and catalase (CAT) activities were all greater in water-stressed seedlings in the 0.2 mg/l brassinolide treatment compared to the control. The results indicate that the application of brassinolide can ameliorate the effects of water stress and enhance drought resistance of Robinia seedlings. Treatment of seedlings with brassinolide may be a useful management tool for afforestation projects in arid and semiarid areas.  相似文献   

12.
The physiological basis of drought resistance in Ziziphus rotundifolia Lamk., which is an important, multipurpose fruit tree of the northwest Indian arid zone, was investigated in a greenhouse experiment. Three irrigation regimes were imposed over a 34-day period: an irrigation treatment, a gradual drought stress treatment (50% of water supplied in the irrigation treatment) and a rapid drought stress treatment (no irrigation). Changes in gas exchange, water relations, carbon isotope composition and solute concentrations of leaves, stems and roots were determined. The differential rate of stress development in the two drought treatments did not result in markedly different physiological responses, but merely affected the time at which they were expressed. The initial response to decreasing soil water content was reduced stomatal conductance, effectively maintaining predawn leaf water potential (Psi(leaf)), controlling water loss and increasing intrinsic water-use efficiency, while optimizing carbon gain during drought. Carbon isotope composition (delta13C) of leaf tissue sap provided a more sensitive indicator of changes in short-term water-use efficiency than delta13C of bulk leaf tissue. As drought developed, osmotic potential at full turgor decreased and total solute concentrations increased in leaves, indicating osmotic adjustment. Decreases in leaf starch concentrations and concomitant increases in hexose sugars and sucrose suggested a shift in carbon partitioning in favor of soluble carbohydrates. In severely drought-stressed leaves, high leaf nitrate reductase activities were paralleled by increases in proline concentration, suggesting an osmoprotective role for proline. As water deficit increased, carbon was remobilized from leaves and preferentially redistributed to stems and roots, and leaves were shed, resulting in reduced whole-plant transpiration and enforced dormancy. Thus, Z. rotundifolia showed a range of responses to different drought intensities indicating a high degree of plasticity in response to water deficits.  相似文献   

13.
Reconstituted dikaryons of Pisolithus sp. (Pers.) Coker & Couch from South Africa influenced growth parameters (shoot length, shoot/root ratio and leaf area), nutrition and physiological indicators (transpiration rate, stomatal conductance and xylem water potential) of maritime pine (Pinus pinaster Ait.) seedlings during drought and recovery from drought. Seedlings colonized with certain dikaryons were more sensitive to water stress and showed less mycorrhiza formation under water stress than seedlings colonized with other dikaryons. Control (uninoculated) seedlings were significantly smaller than those inoculated with dikaryons. Transpiration rate, stomatal conductance and xylem water potential varied among mycorrhizal treatments during the water stress and recovery periods. After rewatering, the controls and seedlings inoculated with dikaryon 34 x 20 had a weaker recovery in transpiration rate, stomatal conductance and xylem water potential than the other treatments and appeared to have experienced damage due to the water stress. Concentrations of various elements differed in the shoots of Pinus pinaster colonized by the various dikaryons. It is suggested that breeding of ectomycorrhizal fungi could constitute a new tool for improving reforestation success in arid and semi-arid zones.  相似文献   

14.
Seedlings of Betula pendula Roth were grown with their root systems separated between two soil compartments. Four treatments were imposed: (i) adequate irrigation in both compartments (WW, controls); (ii) adequate irrigation in one compartment and drought in the other compartment (WD); (iii) drought in both compartments (DD); and (iv) half of the root system severed and the remainder kept well-watered (root excision, RE). Predawn leaf water potential, stomatal conductance, soil-to-leaf specific hydraulic conductance, and root and leaf growth decreased in DD-treated seedlings, which also displayed severe leaf shedding (30% loss in leaf area). The DD treatment also resulted in increased concentrations of abscisic acid (ABA) and its glucose ester in the xylem sap of roots and shoots compared to concentrations in control seedlings (about 200 versus 20 nM). Despite the difference in xylem sap concentrations, total ABA flux to the shoots was similar in the two treatments (1-2 pmol ABA m(-2) leaf area s(-1)) as a result of reduced transpiration in the DD-treated seedlings. Compared with root growth in control plants, root growth increased in the RE-treated plants and decreased in the drying compartment of the WD treatment; however, the RE and WD treatments only slightly reduced leaf expansion, and had no detectable effects on shoot water relations or ABA concentrations of the root and shoot xylem sap. We conclude that short-term soil water depletion affecting only 50% of the root system does not cause a measurable stress response in birch shoots, despite root growth cessation in the fraction of drying soil.  相似文献   

15.
利用中性盐NaCl、碱性盐Na2CO3及混合盐碱(NaCl∶ Na2CO3=1∶1)设置3种盐碱胁迫处理,研究平欧杂种榛“辽榛3号”在不同类型、不同浓度的盐碱胁迫下生长指标及内源激素的变化,分析各内源激素与生长指标的相关关系.结果表明:3种盐碱胁迫对平欧杂种榛株高、新梢长度及叶片数净生长量、地上部生物量、总生物量抑制作用明显,且NaCl胁迫>混合盐碱胁迫>Na2CO3胁迫,而各处理根冠比显著增加;低浓度盐碱胁迫对地径有明显的增粗效应,在50、100 mmol·L-1 Na2CO3胁迫下,地下部生物量显著增加;此外,NaCl胁迫还显著降低了平欧杂种榛叶片含水量.3种胁迫条件下,平欧杂种榛叶片的ABA含量明显增加,且(GA+ IAA+ ZR)/ABA比值与对照相比均显著降低,同时发现Na2CO3胁迫下ABA积累更快,且与胁迫浓度无关,而NaCl胁迫下ABA含量与胁迫浓度显著正相关.相关分析表明:(GA+ IAA+ZR)/ABA比值与株高、新梢长度、新梢叶片数存在显著正相关,而与根冠比呈极显著的负相关.由上可知,平欧杂种榛在中性盐下的生长指标受抑制程度大于碱性盐和混合盐碱,不同盐碱胁迫下内源激素响应特征也存在差异,表明平欧杂种榛对Na2CO3的耐受力高于NaCl.  相似文献   

16.
沙棘抗旱性是多因素作用的结果,如果只利用单因素/单项指标评价其抗逆性,则具有一定的片面性。本研究对干旱胁迫下4个沙棘品种的多项生理生化指标的变化进行比较分析,并通过主成分分析和综合评价发现:乡土中国沙棘(河北丰宁种源)品种的抗旱性最强,引进大果沙棘―乌兰格木品种的抗旱性最差,引进的4个沙棘品种抗旱能力强弱的排序为:丰宁(0.619)> 楚伊(0.445)> 向阳(0.390)> 乌兰格木(0.274)。净光合速率、蒸腾速率、气孔导度、叶水势、谷胱甘肽还原酶和脱落酸的含量可作为沙棘抗旱性的判定指标。随着干旱胁迫的加剧,与对照相比,4个引沙棘品种的净光合速率、蒸腾速率、气孔导度和叶水势均下降,而超氧化物歧化酶活性、细胞膜透性、谷胱甘肽还原酶和四大激素均上升,这说明干旱逆境胁迫影响了沙棘正常的生长发育过程,同时通过内在的生理生化指标的改变来增加其自身对逆境胁迫的抵抗能力。这些将为沙棘抗旱树种的筛选和进一步的杂交育种提供依据。  相似文献   

17.
指出了内源激素作为植物体内的调节者以及痕量信号分子,特别是在植物遭受逆境胁迫时,激素起着重要的调节作用。早期的多数实验表明,植物通过对内源激素的调节,如IAA和CTK浓度的变化以及ABA浓度的升高等内源激素的动态变化来调节某些生理过程以达到适应干旱的效果。目前关于干旱胁迫条件下植物抗氧化代谢的研究较多,而关于其分子以及信号物质调控机制的研究较少,应以内源激素为中间纽带,形成一条分子到生理再到生态,从微观到宏观的植物生理生态的研究链条。  相似文献   

18.
Leaf conductance, water relations, growth, and abscisic acid (ABA) concentrations in xylem sap, root apices and leaves were assessed in oak seedlings (Quercus robur L.) grown with a root system divided between two compartments and subjected to one of four treatments: (a) well watered, WW; (b) half of root system exposed to soil drying and half kept well watered, WD; (c) whole root system exposed to drought, DD; and (d) half of root system severed, RE. Sharp decreases in plant stomatal conductance, leaf water potential, hydraulic conductance and leaf growth were observed during DD treatment. No significant differences in plant leaf water potential and stomatal conductance were detected between the WW and WD treatments. Nevertheless, the WD treatment resulted in inhibition of leaf expansion and stimulation of root elongation only in the well-watered compartment. Abscisic acid concentrations did not change in leaves, root tips, or xylem sap of WD- compared to WW-treated plants. Increased concentrations of ABA were observed in xylem sap from DD-treated plant roots, but the total flux of ABA to shoots was reduced compared to that in WW-treated plants, because of decreases in transpiration flux. Similar plant responses to the WD and RE treatments indicate that the responses observed in the WD-treated plants were probably not triggered by a positive signal originating from drying roots.  相似文献   

19.
Li M  Li Y  Li H  Wu G 《Tree physiology》2011,31(3):349-357
Paper mulberry (Broussonetia papyrifera L. Vent) is well known for its bark fibers, which are used for making paper, cloth, rope, etc. It was found that, in addition to its well-documented role in the enhancement of plant salt tolerance, overexpression of the Na+/H+ antiporter (AtNHX5) gene in paper mulberry plants showed high drought tolerance. After exposure to water deficiency and salt stress, the wild-type (WT) plants all died, while the AtNHX5-overexpressing plants remained alive under high salt stress, and had a higher survival rate (>66%) under drought stress. Measurements of ion levels indicated that Na+ and K+ contents were all higher in AtNHX5-overexpressing leaves than in WT leaves in high saline conditions. The AtNHX5 plants had higher leaf water content and leaf chlorophyll contents, accumulated more proline and soluble sugars, and had less membrane damage than the WT plants under water deficiency and high saline conditions. Taken together, the results indicate that the AtNHX5 gene could enhance the tolerance of paper mulberry plants to multiple environmental stresses by promoting the accumulation of more effective osmolytes (ions, soluble sugars, proline) to counter the osmotic stress caused by abiotic factors.  相似文献   

20.
‘Zhongqiusucui’ jujube secondary shoots were treated with 3-indolebutyric acid (IBA) at three concentrations, 500, 1000 and 1500 mg/L. Results show that IBA could significantly enhance rooting and root characteristics of cuttings and were best with IBA at 1500 mg/L. In the rooting process, the formation of adventitious roots was related to the consumption and accumulation of nutrients (soluble sugars and proteins) and the changes in endogenous hormones in phloem, leaf tips and leaf bases. The rooting of cuttings had a positive correlation with the consumption of soluble sugars during the period of callus formation and with the accumulation of soluble sugars during adventitious root formation and growth. Rooting was positively related to the breakdown of soluble proteins in the phloem when the callus formed, and had a positive correlation with its accumulation during adventitious root formation and growth. Leaf tips and leaf bases showed a reverse trend in changes of soluble protein. However, together with the phloem, leaf tips and leaf bases regulated and controlled the formation and development of adventitious roots. The main activities of soluble proteins exist in the leaf tips as this was the main source of soluble proteins. The relation between rooting and IAA (indole-3-acetic acid) content in phloem was positive and thus a high concentration of IAA could benefit the induction and formation of adventitious roots. However, rooting was negatively related with ABA (abscisic acid) and GA (gibberellic acid) and a high concentration of both could inhibit the induction and formation of adventitious roots. Rooting had a positive correlation with phloem IAA/ABA ratios, and higher ratios could improve rooting. Low concentrations of ZR (zeatin riboside) triggered the induction of adventitious roots, while higher concentrations promoted root growth. Endogenous hormones in leaf tips and bases had an impact on rooting. The activities of endogenous hormones mainly existed in leaf tips because they play a major role in the production and consumption of IAA and its ABA content increased during rooting. The ZR in leaf tips influenced the rooting of cuttings, especially in the callus formation and rooting stage. Leaf tips were the main source of GA.  相似文献   

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