“顶端优势”原理在生产实践中的应用与思考

<正>在新课标人教版高中生物必修3第三章《植物的激素调节》中,以顶端优势为例,说明植物激素之一——生长素对植物生长有两重性——既能促进生长,又能抑制生长。典型体现在植物的"顶端优势"上。顶芽产生生长素由于极性运输从而造成侧芽生长素积累,结果导致顶芽生长快,侧芽生长受到抑制。这在农业生产上对一些农作     

不同供硼水平对绿豆植株形态及其叶片生长特征的影响

利用水培以绿豆为材料,研究不同供硼水平对绿豆植株形态和叶片生长特征的影响。结果表明缺硼抑制绿豆生长,但对根的影响较对冠的影响更大,表现在缺硼导致冠根比增大;缺硼明显抑制叶面积;降低特定叶面积（SLA）,这可能是由于缺硼影响细胞伸展的缘故,造成叶片密度增加,缺硼也提高叶片重量比（LWR)并导致叶脉间失绿,说明缺硼叶片可能过量碳水化合物积累,引起叶绿素降解,与适量供硼比较,过量供硼也影响绿豆的生长,但对冠根比没有影响,表明过量供硼对根和冠具有相同的抑制作用,硼中毒导致成熟叶片脱落,从而影响叶面积,但对特定叶面积（SLA）和叶片重量比（LWR）没有影响。     

强旱生小灌木绵刺劈裂生长过程中内源激素含量变化的研究

 研究强旱生小灌木绵刺(Potaninia mongolica)劈裂生长过程中内源激素含量的变化。结果表明：1）4种生长状态中,完全劈裂的植株的叶片及劈裂发生部位ABA的含量比其它3种状态的都低,而其根中ABA的含量最大。同其它几种激素相比,ABA在绵刺体内的含量最大;2）劈裂生长发生之前,在劈裂发生部位IAA积累量大,尤其是在即将劈裂的过渡植株的劈裂发生部位IAA含量最大;3）劈裂生长发生过程中GA3含量的变化与IAA的变化有同步性;4）ZR的含量也是在劈裂生长发生前的绵刺的劈裂发生部位中较大,随着劈裂生长的发生,植物从根部向叶片及劈裂发生部位运输的ZR有逐渐降低的趋势,而在劈裂生长发生的过渡阶段,ZR从根部向劈裂发生部位运输的比例较大,分别为19.44%和20%;5）IAA、GA3、ZR 三者协调促进劈裂发生部位细胞的生长和分裂,而ABA的积累对绵刺适应干旱的环境条件起到了一定的调节作用。     

多效唑调节水稻植株生长的作用机理

多效唑处理水稻植株后使株高降低和分蘖增加,植株体内的GA类物质和IAA含量相应下降,乙烯释放率显著增加;用外源GA_3和氮肥能逆转多效唑的抑制作用而使株高增加,同时GA类物质含量也明显增加。因此,多效唑不仅调节体内的GA类物质含量,还能通过调节IAA含量和乙烯释放率而影响植物生长。     

油菜幼苗对硼的吸收与运转及钙的影响

硼促进油菜幼苗生长和对N、P、K、Ca 营养元素的吸收.当油菜生长介质中钙浓度增大时,植株吸硼量和硼在新叶中的积累减少,老叶中积累增加。硼在油菜体内的运转系数在低硼条件下较大,并随钙处理水平的增加而下降.结果表明,在硼素供应有限的条件下,油菜体内硼仍具一定的移动性,钙则使硼的吸收和移动性下降.     

氮碳源对冬凌草再生植株生长及次生代谢产物的影响

以冬凌草再生植株为研究材料,通过改变培养基中蔗糖浓度及NO3-/NH4+比例,研究冬凌草再生植株的生长状况及植株体内次生代谢物的积累规律.结果表明:3%蔗糖有利于冬凌草再生植株的生长和冬凌草甲素的积累,5%蔗糖有利于迷迭香酸的积累;NO3-/NH4+比例过高或过低均不利于再生植株的生长和次生代谢产物的积累,二者比例为2∶1时再生植株生长和次生代谢产物积累最佳.说明不同蔗糖浓度及NO3-/NH4+比例对冬凌草再生植株的生长和次生代谢物合成有明显的影响.     

生长素极性运输的自动抑制

以豌豆(Pisum sativum L.)和绿豆(Phaseolus radiatus L.)为材料证明了IAA极性运输的自动抑制现象。用改进的“供体-受体技术”证明:受体中的IAAA可抑制~3H-IAA的极性运输,抑制程度随受体中加入IAA浓度的增加而增强;在“Y”型外植体一侧切口施用的IAA也可抑制~3H-IAA在另一侧的极性运输,并导致~3H-IAA在该侧组织中的积累以及代谢或钝化的增强。此外,组织中游离~3H-IAA的比例也随运输时间的延长而不断减小。     

新疆胀果甘草幼苗耐盐性及对NaCl胁迫的离子响应

以盐碱荒漠草甸药用植物胀果甘草(Glycyrrhiza inflata)为材料, 采用水培法研究了盐处理(50、100、200、300 mmol·L^-1NaCl) 28天后幼苗株高、生物量、含水量、根粗、甘草酸含量和不同器官的离子含量及离子的选择吸收、运输能力, 并对丙二醛、脯氨酸含量进行测定, 以确定其耐盐范围及耐盐方式。结果表明, 低盐浓度对胀果甘草幼苗生长无显著影响, 只有较高盐浓度(≥200 mmol·L^-1 NaCl)使幼苗总生物量、株高、甘草酸含量显著降低; 根据耐盐系数与盐浓度的拟合方程, 确定适宜幼苗生长的盐浓度范围为0-278.17 mmol·L^-1。随盐浓度上升, 植株选择性吸收K⁺、Ca²⁺、Mg²⁺, 而抑制Na⁺进入体内, 幼苗对进入植株体内的Na⁺在不同盐浓度下采取了不同的分配策略, 低盐浓度下(0-100 mmol·L^-1), 植株体内Na⁺主要积累在根中, 避免了叶中Na⁺的过多积累, 其盐适应机制以耐盐方式为主; 高盐浓度下(≥200 mmol·L^-1 NaCl), Na⁺主要积累在下部叶, 并通过叶片脱落的方式带走体内的盐分, 其盐适应机制以避盐方式为主。盐胁迫下, 幼苗能促进K⁺而抑制Na⁺向上部叶的运输, 使上部叶拒Na喜K, 维持了较高的K⁺/Na⁺比值, 有利于幼苗生长; 同时, 地下根系能通过积累Ca²⁺、Mg²⁺和合成脯氨酸、甘草酸, 以提高渗透调节能力, 缓解Na⁺毒害, 使根的生长不受影响, 有利于保证幼苗在盐环境中吸收维持生长的必要养分, 这是胀果甘草幼苗具有较强耐盐性的原因。以上结果说明, 胀果甘草幼苗通过对盐离子的吸收和运输调控、离子区域化和渗透调节, 以耐盐和避盐两种方式适应盐碱荒漠环境。     

乙烯在豌豆植株顶端优势中的作用

用人工合成细胞分裂素BAP和CPPU处理豌豆植株叶腋可诱导处理部位侧芽的生长,同时伴有大量乙烯产生;用乙烯合成抑制剂AVG处理或植株去顶同样可导致创芽生长,但乙烯释放量却明显少于对照,表明侧芽的生长与乙烯释放量的多少无关。而3种物质处理后诱导产生的侧芽的数目、长度及其鲜重与处理部位内源IAA含量的增加则呈正相关。     

缺硼与低温对黄瓜幼苗—些生理反应的影响

营养液培养试验表明,缺硼明显降低黄瓜幼苗植株的生长量,植株叶片细胞质膜K 的渗漏增加,细胞结构受损。此外,叶片中蔗糖和还原糖都明显积累。缺硼的上述反应在低温胁迫（7－8℃,FX25－28℃为对照）时表现更为明显。试验结果表明硼对于细胞膜的完整性具有重要的作用;充分供硼可以减轻低温对细胞膜的伤害。     

Auxin dynamics after decapitation are not correlated with the initial growth of axillary buds

One of the first and most enduring roles identified for the plant hormone auxin is the mediation of apical dominance. Many reports have claimed that reduced stem indole-3-acetic acid (IAA) levels and/or reduced basipetal IAA transport directly or indirectly initiate bud growth in decapitated plants. We have tested whether auxin inhibits the initial stage of bud release, or subsequent stages, in garden pea (Pisum sativum) by providing a rigorous examination of the dynamics of auxin level, auxin transport, and axillary bud growth. We demonstrate that after decapitation, initial bud growth occurs prior to changes in IAA level or transport in surrounding stem tissue and is not prevented by an acropetal supply of exogenous auxin. We also show that auxin transport inhibitors cause a similar auxin depletion as decapitation, but do not stimulate bud growth within our experimental time-frame. These results indicate that decapitation may trigger initial bud growth via an auxin-independent mechanism. We propose that auxin operates after this initial stage, mediating apical dominance via autoregulation of buds that are already in transition toward sustained growth.     

Response to strigolactone treatment in chrysanthemum axillary buds is influenced by auxin transport inhibition and sucrose availability

Axillary bud outgrowth is regulated by both environmental cues and internal plant hormone signaling. Central to this regulation is the balance between auxins, cytokinins, and strigolactones. Auxins are transported basipetally and inhibit the axillary bud outgrowth indirectly by either restricting auxin export from the axillary buds to the stem (canalization model) or inducing strigolactone biosynthesis and limiting cytokinin levels (second messenger model). Both models have supporting evidence and are not mutually exclusive. In this study, we used a modified split-plate bioassay to apply different plant growth regulators to isolated stem segments of chrysanthemum and measure their effect on axillary bud growth. Results showed axillary bud outgrowth in the bioassay within 5 days after nodal stem excision. Treatments with apical auxin (IAA) inhibited bud outgrowth which was counteracted by treatments with basal cytokinins (TDZ, zeatin, 2-ip). Treatments with basal strigolactone (GR24) could inhibit axillary bud growth without an apical auxin treatment. GR24 inhibition of axillary buds could be counteracted with auxin transport inhibitors (TIBA and NPA). Treatments with sucrose in the medium resulted in stronger axillary bud growth, which could be inhibited with apical auxin treatment but not with basal strigolactone treatment. These observations provide support for both the canalization model and the second messenger model with, on the one hand, the influence of auxin transport on strigolactone inhibition of axillary buds and, on the other hand, the inhibition of axillary bud growth by strigolactone without an apical auxin source. The inability of GR24 to inhibit bud growth in a sucrose treatment raises an interesting question about the role of strigolactone and sucrose in axillary bud outgrowth and calls for further investigation.     

Involvement of auxin and CKs in boron deficiency induced changes in apical dominance of pea plants (Pisum sativum L.)

It has previously been shown that boron (B) deficiency inhibits growth of the plant apex, which consequently results in a relatively weak apical dominance, and a subsequent sprouting of lateral buds. Auxin and cytokinins (CKs) are the two most important phytohormones involved in the regulation of apical dominance. In this study, the possible involvement of these two hormones in B-deficiency-induced changes in apical dominance was investigated by applying B or the synthetic CK CPPU to the shoot apex of pea plants grown in nutrient solution without B supply. Export of IAA out of the shoot apex, as well as the level of IAA, Z/ZR and isopentenyl-adenine/isopentenyl-adenosine (i-Ade/i-Ado) in the shoot apex were assayed. In addition, polar IAA transport capacity was measured in two internodes of different ages using 3H-IAA. In B-deficient plants, both the level of auxin and CKs were reduced, and the export of auxin from the shoot apex was considerably decreased relative to plants well supplied with B. Application of B to the shoot apex restored the endogenous Z/ZR and IAA level to control levels and increased the export of IAA from the shoot apex, as well as the 3H-IAA transport capacity in the newly developed internodes. Further, B application to the shoot apex inhibited lateral bud growth and stimulated lateral root formation, presumably by stimulated polar IAA transport. Applying CPPU to the shoot apex, a treatment that stimulates IAA export under adequate B supply, considerably reduced the endogenous Z/ZR concentration in the shoot apex, but had no stimulatory effect on IAA concentration and transport in B-deficient plants. A similar situation appeared to exist in lateral buds of B-deficient plants as, in contrast to plants well supplied with B, application of CKs to these plants did not stimulate lateral bud growth. In contrast to the changes of Z/ZR levels in the shoot apex, which occurred after application of B or CPPU, the levels of i-Ade/i-Ado stayed more or less constant. These results suggest that there is a complex interaction between B supply and plant hormones, with a B-deficiency-induced inhibition of IAA export from the shoot apex as one of the earliest measurable events.     

Modalités du transport et du métabolisme de l'AIA 14C ou AIA 3H en relation avec la morphogenèse des bourgeons axillaires chez le Scrophularia arguta

Modes of transport and metabolism of ¹⁴ C-IAA and ³ H-IAA in relation to morphogenesis of axillary buds in Scrophularia arguta. The main objectives of this study were to investigate the morphogenetic role of IAA on the growth and development of axillary buds. After foliar applications of radioactive IAA for 6 h on intact plants of Scrophularia arguta Sol. the characteristics of auxin transport were studied by liquid scintillation counting, thin layer chromatography and microautoradiography. The main part of the radioactivity moved at a mean rate of 7 mm/h. Over long periods of transport, the tracers accumulated at the base of the axis and in the roots. The nodes were a little richer in ³H or ¹⁴C than the internodes. This fact seemed to be correlated with the vascular organization of this part of the stem. A very weak proportion of tracers was found in axillary buds. The radioactivity was to about 50% associated with the IAA molecule; the rest corresponded essentially to indolyl-4-acetyl-l-aspartic acid and indolyl-3-aldehyde. Tracers were mainly concentrated in the phloem along the whole axis and, to a lesser extent in some of the young differentiating metaxylem vessels, and in the medullary rays. No radioactivity was found in the cambial zone and in the mature xylem, nor in the parenchymas. These results support the view of an indirect role of IAA on the axillary bud growth and morphogenesis.     

Origin and basipetal transport of the IAA responsible for rooting of carnation cuttings

The origin and transport of the IAA responsible for rooting was studied in carnation (Dianthus caryophyllus L.) cuttings obtained from secondary shoots of the mother plants. The presence of mature leaves in the cuttings was essential for rooting. Removal of the apex and/or the youngest leaves did not reduce the rooting percentage as long as mature leaves remained attached. Removal of mature leaves inhibited rooting for a 24-day period during which the basal leaves grew and reached maturity. After this period rooting progressed as in intact cuttings. Auxin (NAA + IBA) applied to the stem base of defoliated cuttings was about 60% as effective as mature leaves in stimulating rooting. Application of NPA to the basal internode resulted in full inhibition of rooting. The view, deduced from these results, that auxin from mature leaves is the main factor controlling the rooting process was reinforced by the fact that mature leaves contained IAA and exported labelled IAA to the stem. The distribution of radioactivity after application of (5-3H)-IAA to mature leaves showed that auxin movement in the stem was basipetal and sensitive to NPA inhibition. The features of this transport were studied by applying 3H-IAA to the apical cut surface of stem sections excised from cuttings. The intensity of the transport was lower in the oldest node than in the basal internode, probably due to the presence of vascular traces of leaves. Irrespective of the localization of the sections and the carnation cultivar used, basipetal IAA transport was severely reduced when the temperature was lowered from 25 to 4 degrees C. The polar nature of the IAA transport in the sections was confirmed by the inhibition produced by NPA. Local application of IAA to different tissues of the sections revealed that polar auxin transport was associated with the vascular cylinder, the transport in the pith and cortex being low and apolar. The present results strongly support the conclusion that IAA originating from the leaves and transported in the stem through the polar auxin transport pathway was decisive in controlling adventitious rooting.     

Patterns of auxin and abscisic acid movement in the tips of gravistimulated primary roots of maize

Because both abscisic acid (ABA) and auxin (IAA) have been suggested as possible chemical mediators of differential growth during root gravitropism, we compared with redistribution of label from applied ³H-IAA and ³H-ABA during maize root gravitropism and examined the relative basipetal movement of ³H-IAA and ³H-ABA applied to the caps of vertical roots. Lateral movement of ³H-ABA across the tips of vertical roots was non-polar and about 2-fold greater than lateral movement of ³H-IAA (also non-polar). The greater movement of ABA was not due to enhanced uptake since the uptake of ³H-IAA was greater than that of ³H-ABA. Basipetal movement of label from ³H-IAA or ³H-ABA applied to the root cap was determined by measuring radioactivity in successive 1 mm sections behind the tip 90 minutes after application. ABA remained largely in the first mm (point of application) whereas IAA was concentrated in the region 2–4 mm from the tip with substantial levels found 7–8 mm from the tip. Pretreatment with inhibitors of polar auxin transport decreased both gravicurvature and the basipetal movement of IAA. When roots were placed horizontally, the movement of ³H-IAA from top to bottom across the cap was enhanced relative to movement from bottom to top whereas the pattern of movement of label from ³H-ABA was unaffected. These results are consistent with the hypothesis that IAA plays a role in root gravitropism but contrary to the idea that gravi-induced asymmetric distribution of ABA contributes to the response.     

The transport of radiolabeled indoleacetic acid and its conjugates in nodal stem segments of Phaseolus vulgaris L.

The transport of radiolabeled indoleacetic acid (IAA), and some of its conjugates, was investigated in nodal stem segments of Phaseolus vulgaris L. Donor agar blocks containing either [2-acetyl-¹⁴C]-IAA; [2-acetyl-¹⁴C]-indole-3-acetyl-L-aspartate (IAAsp); [2-acetyl-¹⁴C]-indole-3-acetyl-L-glycine (IAGly); or [2-acetyl-¹⁴C]-indole-3-acetyl-L-alanine (IAAla) were placed on either the apical or basal cut surface of stem segments each bearing an axillary bud at the midline. In some experiments, a receiver block was placed on the end opposite to the donor. After transport was terminated, the segments were divided into five equal sections plus the bud, and the radioactivity of donors, receivers and each part of the stem segment was counted.For all four substances tested, the amount of ¹⁴C transported to the axillary bud from the base was the same or greater than that from the apical end. After basipetal transport, the distribution of ¹⁴C in the segment declined sharply from apex to base. The inverse was true for acropetal transport. Transport for the three IAA conjugates did not differ substantially from each other.The IAA transport inhibitor, N-1-naphthylphthalamic acid (NPA), inhibited basipetal ¹⁴C-IAA transport to the base of the stem segment but did not alter substantially the amount of ¹⁴C-IAA recovered from the bud. Transport of ¹⁴C-IAA from the apical end to all parts of the stem segment declined when the base of the section was treated with nonradioactive IAA. Taken together with data presented in the accompanying article [Tamas et al. (1989) Plant Growth Regul 8: 165–183], these results suggest that the transport of IAA plays a role in axillary bud growth regulation, but its effect does not depend on the accumulation of IAA in the axillary bud itself.     

植物生长调节剂对黑木相思优树腋芽增殖及生根的影响

为建立黑木相思(Acacia melanoxylon)快繁技术体系,以含1个腋芽的无菌茎段为材料,研究了植物生长调节剂对其增殖和生根的影响。结果表明,6-BA 极易诱导黑木相思愈伤组织形成,但芽长势较差,不利于腋芽增殖体系的建立。而生长素既能诱导黑木相思生根,又能诱导腋芽增殖;将无菌茎段接入MS+IAA 0.5 mg L^-1+IBA 0.5 mg L^-1培养基中培养20 d的生根率为98.41%,培养40 d的腋芽增殖倍数为2.36,单株繁殖系数为6.57。这是首次成功建立高效、简便的生根和增殖同步发生的黑木相思直接器官发生途径的组培技术体系。     

Influence of 2,3,5-Triiodobenzoic Acid and 1-N-Naphthylphthalamic Acid on Indoleacetic Acid Transport in Carnation Cuttings: Relationship with Rooting

³H-IAA transport in excised sections of carnation cuttings was studied by using two receiver systems for recovery of transported radioactivity: agar blocks (A) and wells containing a buffer solution (B). When receivers were periodically renewed, transport continued for up to 8 h and ceased before 24 h. If receivers were not renewed, IAA transport decreased drastically due to immobilization in the base of the sections. TIBA was as effective as NPA in inhibiting the basipetal transport irrespective of the application site (the basal or the apical side of sections). The polarity of IAA transport was determined by measuring the polar ratio (basipetal/acropetal) and the inhibition caused by TIBA or NPA. The polar ratio varied with receiver, whereas the inhibition by TIBA or NPA was similar. Distribution of immobilized radioactivity along the sections after a transport period of 24 h showed that the application of TIBA to the apical side or NPA to the basal side of sections, increased the radioactivity in zones further from the application site, which agrees with a basipetal and acropetal movement of TIBA and NPA, respectively. The existence of a slow acropetal movement of the inhibitor was confirmed by using ³H-NPA. From the results obtained, a methodological approach is proposed to measure the variations in polar auxin transport. This method was used to investigate whether the variations in rooting observed during the cold storage of cuttings might be related to changes in polar auxin transport. As the storage period increased, a decrease in intensity and polarity of auxin transport occurred, which was accompanied by a delay in the formation and growth of adventitious roots, confirming the involvement of polar auxin transport in supplying the auxin for rooting. Received April 19, 1999; accepted December 2, 1999     

Effect of plant growth substances on the growth of axillary buds in cultured stem segments of Phaseolus vulgaris L.

The hormonal control of axillary bud growth was investigated in cultured stem segments of Phaseolus vulgaris L. When the stem explants were excised and implanted with their apical end in a solid nutrient medium, outgrowth of the axillary buds-located at the midline of the segment-was induced. However, if indoleacetic acid (IAA) or naphthaleneacetic acid (NAA) was included in the medium, bud growth was inhibited. The exposure of the apical end to IAA also caused bud abscission and prevented the appearance of new lateral buds.In contrast to apically inserted segments, those implanted in the control medium with their basal end showed much less bud growth. In these segments, the auxin added to the medium either had no effect or caused a slight stimulation of bud growth.The IAA transport inhibitor N-1-naphthylphthalamic acid (NPA) relieved bud growth inhibition by IAA. This suggests that the effect of IAA applied at the apical end requires the transport of IAA itself rather than a second factor. With the apical end of the segment inserted into the IAA-containing medium, simultaneous basal application of IAA relieved to some extent the inhibitory effect of the apical IAA treatment. These results, together with data presented in a related article [Lim R and Tamas I (1989) Plant Growth Regul 8: 151–164], show that the polarity of IAA transport is a critical factor in the control of axillary bud growth.Of the IAA conjugates tested for their effect on axillary bud growth, indoleacetyl alanine, indoleacetic acid ethyl ester, indoleacetyl-myo-inositol and indoleacetyl glucopyranose were strongly inhibitory when they were applied to the apical end of the stem explants. There was a modest reduction of growth by indoleacetyl glycine and indoleacetyl phenylalanine. Indoleacetyl aspartic acid and indoleglyoxylic acid had no effect.In addition to IAA and its conjugates, a number of other plant growth substances also affected axillary bud growth when applied to the apical end of stem segments. Myo-inositol caused some increase in the rate of growth, but it slightly enhanced the inhibitory effect of IAA when the two substances were added together. Gibberellic acid (GA₃) caused some stimulation of bud growth when the explants were from younger, rather than older plants. The presence of abscisic acid (ABA) in the medium had no effect on axillary bud growth. Both kinetin and zeatin caused some inhibition of axillary buds from younger plants but had the opposite effect on buds from older ones. Kinetin also enhanced the inhibitory effect of IAA when the two were applied together.In conclusion, axillary buds of cultured stem segments showed great sensitivity to auxins and certain other substances. Their growth responded to polarity effects and the interaction among different substances. Therefore, the use of cultured stem segments seems to offer a convenient, sensitive and versatile test system for the study of axillary bud growth regulation.