The effect of simulated microgravity on seed germination and seedling anatomy of Phaseolus vulgaris L

Seed germination and root anatomy were investigated in seedlings of Phaseolus vulgaris L. developed on a slowly rotating bi-dimensional clinostat and in 1g. Germination time, percent germination, curvature and anatomy of developing root apexes were monitored on the clinostat and compared with the control. Interesting differences were found in germination and root features of the seeds developed on the clinostat compared with 1g ones: the main being germination time, root cap formation, the quantity and distribution of amyloplasts in statocytes. The use of a software to quantitatively analyse root cap anatomy allowed us to detect some differences otherwise unlikely to highlight. Our results showed that prolonged rotation on a bi-dimensional clinostat has an effect on some aspects of germination and on the statocytes that continuously perceives gravity from ever-changing directions.     

Growth and Development in Arabidopsis thaliana through an Entire Life Cycle under Simulated Microgravity Conditions on a Clinostat

Arabidopsis thaliana ecotype Columbia and Landsberg erecta were studied. Horizontal clinorotation affected little germination of seeds, growth and development of rosette leaves and roots during early vegetative growth stage, and the onset of the bolting of inflorescence axis and flower formation in reproductive growth stage, although it suppressed elongation of inflorescence axes. The clinorotation substantially reduced the numbers of siliques and seeds in Landsberg erecta, and completely inhibited seed production in Columbia. Seeds produced in Landsberg erecta on the clinostat were capable of germinating and developing rosette leaves normally on the ground. On the other hand, growth of pin-formed mutant (pin/pin) of Arabidopsis ecotype Enkheim, which has a unique structure of inflorescence axis with no flower and extremely low levels of auxin polar transport activity, was inhibited and the seedlings frequently died during vegetative stage on the clinostat. Seed formation and inflorescence growth of the seedlings with normal shape (pin/+ or +/+) were also suppressed on the clinostat. These results suggest that the growth and development of Arabidopsis, especially in reproductive growth stage, is suppressed under simulated microgravity conditions on a clinostat. To complete the life cycle probably seems to be quite difficult, although it is possible in some ecotypes. Received 18 June 1999/ Accepted in revised form 27 August 1999     

STUDIES ON THE GERMINATION OF SEEDS OF THE ROOT PARASITES,ALECTRA VOGELII AND STRIGA GESNERIOIDES. I. ANATOMICAL CHANGES IN THE EMBRYOS

Anatomical changes in the radicle and shoot meristems of embryos of germinating seeds of the obligate root parasites, Alectra vogelii and Striga gesnerioides were studied. When germination of seeds was stimulated by cowpea (Vigna unguiculata) root exudate, growth occurred mainly in the radicular pole of embryos and minimally in the plumular pole, resulting in seedlings with elongated radicles. Maximum radicle elongations of about 3 mm in A. vogelii and 2 mm in S. gesnerioides were recorded during a period of 8 and 11 days, respectively. Analysis of the radicular tip during the course of seed germination revealed that the activity of the meristematic tissue progressively decreased until it completely disappeared. When germinated seeds were cultured on nutrient agar media, the radicle meristem of A. vogelii continued to grow producing a normal root with a root cap. On the other hand, the radicles of cultured S. gesnerioides seeds elongated only slightly before meristematic activity ceased. During continued growth of seedlings of both species on agar media, lateral roots whose tips had typical angiosperm root topography, were initiated from the radicle.     

Restitution of polarity in statocytes from centrifuged roots*

Abstract The structural polarity of statocytes from cress roots is changed by centrifugation. Upon low- dose centrifugation (3000 g min), the extent of stratification depends on statocyte position, i.e., central statocytes are affected more than lateral ones. Upon higher doses of centrifugation (60,000 and 360,000 g min), a uniform density gradient is established in all statocytes. If, after centrifugation, the roots are exposed to gravity again, the endoplasmic reticulum (ER) cisternae are relocated parallel to the periclinal cell walls within a few minutes; this relocation is independent of the direction of gravity in relation to the root axis, and independent of the previously applied centrifugation dose. This supports the notion that polarity is determined genetically. Cytochalasin B treatment, before and during centrifugation, totally inhibits the relocation of ER. After removing the drug by rinsing the roots, the statocytes restore cell polarity and relocate ER. These results indicate that relocation of ER cisternae may be mediated by microfilaments. When centrifuged roots are exposed to 1 g in the horizontal position, the latent period of gravitropism increases by 8–10 min relative to controls, regardless of the previously applied centrifugation doses. The kinetics of curvature are virtually identical. Since the increase in the latent period coincides with the time needed for most statocytes to restore the distal cell pole, it is evident that perception of gravity is correlated to the integrity of the distal cell pole.     

Enhancement of phototropic response to a range of light doses in Triticum aestivum coleoptiles in clinostat-simulated microgravity

The phototropic dose-response relationship has been determined for Triticum aestivum cv. Broom coleoptiles growing on a purpose-built clinostat apparatus providing gravity compensation by rotation about a horizontal axis at 2 rev·min^-1. These data are compared with data sets obtained with the clinostat axis vertical and stationary, as a 1·g control, and rotating vertically to examine clinostat effects other than gravity compensation. Triticum at 1·g follows the wellestablished pattern of other cereal coleoptiles with a first positive curvature at low doses, followed by an indifferent response region, and a second positive response at progressively increasing doses. However, these response regions lie at higher dose levels than reported for Avena. There is no significant difference between the responses observed with the clinostat axis vertical in the rotating and stationary modes, but gravity compensation by horizontal rotation increases the magnitude of first and second positive curvatures some threefold at 100 min after stimulation. The indifferent response is replaced by a significant curvature towards the light source, but remains apparent as a reduced curvature response at these dose levels.     

Gravitropic bending of cress roots without contact between amyloplasts and complexes of endoplasmic reticulum

The polar arrangement of cell organelles in Lepidium root statocytes is persistently converted to a physical stratification during lateral centrifugation (the centrifugal force acts perpendicular to the root long axis) or by apically directed centrifugation combined with cytochalasin-treatment. Lateral centrifugation (10 min, 60 min at 10\g or 50\g) causes displacement of amylplasts to the centrifugal anticlinal cell wall and shifting of the endoplasmic reticulum (ER) complex to the centripetal distal cell edge. After 60 min of lateral centrifugation at 10\g or 50\g all roots show a clear gravitropic curvature. The average angle of curvature is about 40° and corresponds to that of roots stimulated gravitropically in the horizontal position at 1\g in spite of the fact that the gravistimulus is 10-or 50-fold higher. Apically directed centrifugation combined with cytochalasin B (25 g\ml^-1) or cytochalasin D (2.5 g\ml^-1) incubation yields statocytes with the amyloplasts sedimented close to the centrifugal periclinal cell wall and ER cisternae accumulated at the proximal cell pole. Gravitropic stimulation for 30 min in the horizontal position at 1\g and additional 3 h rotation on a clinostat result in gravicurvature of cytochalasin B-treated centrifuged (1 h at 50\g) roots, but because of retarded root growth the angle of curvature is lower than in control roots. Cytochalasin D-treatment during centrifugation (20 min at 50\g) does not affect either root growth or gravicurvature during 3 h horizontal exposure to 1\g relative to untreated roots. As lateral centrifugation enables only short-term contact between the amyloplasts and the distal ER complex at the onset of centrifugation and apically directed centrifugation combined with cytochalasin-treatment even exclude any contact the integrity of the distal cell pole need not necessarily be a prerequisite for graviperception in Lepidium root statocytes.Abbreviations CB cytochalasin B - CD cytochalasin D - ER endoplasmic reticulum - g gravitational acceleration     

Morphogenesis of the protocorm ofCypripedium acaule (Orchidaceae)

The nature of the protocorm of theOrchidaceae has fascinated morphologists for more than a century. In the present study, the development of the protocorm was followed using in vitro germination of seeds on a culture medium containing sugar, but without a symbiont. Inside the seed, the embryo consists of about a hundred cells. In the embryo, cells are arranged along a longitudinal axis according to size; these cells contain protein and lipid reserve material. In the first stages of seedling development, the embryo is transformed into a protocorm and meristematic tissue becomes organized into a meristematic dome (promeristem) at the anterior pole. This meristematic dome will give rise to a scale and the apex of the seedling. At first, the apex and the scale leaf develop synchronously. The development of the root always follows that of the apex. The study of the development of the seed ofCypripedium acaule showed that the protocorm is a distinct morphological system with respect to the rest of the cormus. The protocorm may be interpreted as an extension of the proembryonic stage.     

西双版纳地区4种榕树果实的资源分配及其种子萌发特性

为了解榕树果实的资源分配和种子萌发特征,对西双版纳地区的垂叶榕(Ficus benjamina)、高山榕(F.altissima)、聚果榕(F.racemosa)和无柄雅榕(F.concinna)的果实营养、繁殖资源分配,以及种子萌发特性进行了研究。结果表明,4种榕果的营养分配存在差异,榕果重的种类果肉亦多,榕果大者果腔较大,且种子数量较多,但雄花和榕小蜂数量不一定多。榕果分配给雌性功能及雄性功能繁殖的小花比例也不相同,聚果榕在雌性功能上的分配显著大于雄性功能,其它3种榕果则分配于雄性功能的资源比雌性功能的多,但程度不同。此外,4种榕树的种子大小及重量存在显著的种间差异,以至于影响到种子的萌发率及根茎生长,果实最大的聚果榕种子最多,萌发率高,但由于种子最小,其根茎生长缓慢且短;果实适中的垂叶榕和高山榕种子较大,不但萌发率较高,其根茎生长也较快;果实和种子都小的无柄雅榕,种子萌发率低,根茎生长慢。生长在水热环境好的榕树比生长在相对贫瘠环境的榕树,由于极少面临水热光照等限制,更容易进化出较小的种子,萌发能力和速度也相对较差。这说明不同生境中榕树进化出了不同的繁殖分配机制及资源利用策略来适应环境变化。     

外源水杨酸对盐胁迫下大黄种子萌发及幼苗生长的影响

该研究以掌叶大黄、唐古特大黄和药用大黄种子为材料,采用双层滤纸培养法,设置系列浓度NaCl (0、100、150、200、250 mmol/L) 胁迫试验,以及系列浓度水杨酸(SA)溶液(0、50、100、150、200、250 mg/L)拌种和浸种后盐胁迫实验,测定3种大黄种子萌发及幼苗生长指标,揭示外源水杨酸对盐胁迫下大黄种子萌发及幼苗生长的影响。结果显示：(1)随NaCl浓度增大3种大黄种子的发芽率均呈直线下降趋势,且子叶、胚轴、根和苗等生长均受到强烈抑制。(2)在拌种条件下, 200 mmol/L NaCl胁迫下掌叶大黄苗长在200 mg/L SA处理下受到显著促进; 200 mmol/L NaCl浓度盐胁迫下唐古特大黄种子发芽率在250 mg/L SA处理下受到显著抑制;100 mmol/L NaCl胁迫下药用大黄种子发芽势在200 mg/L SA处理下受到显著抑制,其发芽率在150 mg/L SA处理下得到显著抑制,其苗长在250 mg/L SA处理下受到显著抑制。(3)在浸种条件下, 200 mmol/L NaCl胁迫下掌叶大黄种子发芽率在50 mg/L SA处理下显著提高,其幼苗根长和苗长的生长在250 mg/L SA处理受到显著促进;200 mmol/L NaCl胁迫下唐古特大黄种子的发芽势在200 mg/L SA处理下得到显著促进,其幼苗根和苗的生长在50 mg/L SA处理下得到显著促进;100 mmol/L NaCl 胁迫下药用大黄根和苗的生长在100 mg/L SA处理下均得到显著促进。研究表明,3种大黄种子和幼苗对盐胁迫的响应趋势一致,但对不同浓度SA拌种和浸种的响应有较大差异。     

Ontogeny of the mycoheterotrophic species Afrothismia hydra (Burmanniaceae)

The complete ontogeny of the mycoheterotrophic Afrothismia hydra (Burmanniaceae) from seed to seed dispersal is presented. The oblong–ovoidal seeds are up to 0.7 mm long. They germinate with root tissue only, disrupting the seed coat and developing a primary ovoid root tubercle. At the proximal end of the tubercle, a second tubercle arises and further root initials indicate the sequential growth of more root tubercles with filiform extensions resulting in a small root aggregate. The seed coat often remains attached to this structure. When the root aggregate enlarges, a central axis to which all roots are connected becomes visible. This axis has a growth pole where new root tubercles arise. The same growth pole will later develop into a stem with scale leaves finally terminating in a flower. Flowers develop sympodially when the mature plant is only several centimetres long. After anthesis, the corolla tube disintegrates, leaving a pyxidium which opens by means of a peculiar elongating placenta, here called 'placentophore'. The placentophore elevates the placenta with attached seeds above the flowering level and is interpreted as an adaptation to ombrohydrochory. The reduction of hypocotyl, cotyledon and primary shoot is discussed with regard to the classical germination concepts of monocotyledons and with mycoheterotrophic dicotyledons.  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 31–36.     

Induction of gravity-dependent plasmatic responses in root statocytes by short time contact between amyloplasts and the distal endoplasmic reticulum complex

Statocytes from roots of Lepidium sativum L., which developed after a 2-min soaking on a horizontal clinostat (2 rotations per min) for 44 h, exhibit the same polarity as in vertically grown roots, as indicated by a complex of endoplasmic reticulum (ER) cisternae at the distal cell pole. Amyloplasts are distributed randomly. The kinetics of graviresponse (=curvature) of such roots are identical to those of normally grown roots. Ten-minute exposure of the root, after 24 h development on the clinostat with gravity acting towards the root's basis (inversion), induces no changes in statocyte ultrastructure. However, corresponding exposure in normal orientation leads to subsequent disintegration of the distal ER complex, loss of amyloplast starch, confluence of lipid droplets, and an increase of the lytic compartment. These ultrastructural events thus appear to be induced by a physical contact — however short — between amyloplasts and the distal ER complex.Abbreviation ER endoplasmic reticulum Dedicated to Professor Dr. W. Haupt on the occasion of his 60th birthday     

Cell cycle and cell differentiation in lentil roots grown on a slowly rotating clinostat

Root growth was studied for seedlings of lentil ( Lens culinaris L. cv. Verte du Puy) grown for 27 h either on a slowly rotating clinostat (0.9 rev. min⁻¹) of vertically (controls). Horizontal clinorotation was employed, so that the longitudinal axis of the root was parallel to the axis of the rotation. Morphological (root length and orientation) and cellular (cell proliferation and cell elongation) parameters were studied. The cell cycle was also analysed by flow cytometry. Root length deviation of the roots from the initial orientation was observed on the clinostat; this deviation could be due to spontaneous oscillation. Cell elongation of the clinostat-rotated roots occurred closer to the tip than in the vertical roots, but the mitotic index was not modified. Clinorotation did not change the frequencies of the G1, S and G2 phases of the cell cycle. These results were compared to those obtained during the D1 mission on Spacelab, 1985. The effects of microgravity on root orientation and mitotic index were not simulated by clinorotation.     

A role for fruit structure in seed survival and germination of Swartzia langsdorffii Raddi beyond dispersal

• Diaspore structure has been hypothesised to play a role in seed viability and/or germination of recalcitrant seeds, especially for Swartzia langsdorffii. Thus, this work aims to (i) investigate the in situ contribution of pericarp and aril on seed viability and germination, and (ii) identify morphoanatomical traits of S. langsdorffii diaspores that allow its desiccation‐sensitive seeds to remain viable.
• The role of the pericarp and aril in seed survival and germination was investigated by placing the whole fruit, whole seeds (arillate seed) and bare seeds (without aril) in soil in the forest understorey, assessing germination, emergence, dead, firm and predated seeds, and water content of pericarps, arils and seeds. Correlation analysis was performed between environmental variables and physiological parameters. Histochemical features of diaspores were also investigated.
• Pericarp water content fell after several months, while the aril maintained its water content. Seeds did not lose water even without the presence of the pericarp and aril. However, presence of the pericarp promoted seed water content, viability and germination long after dispersal. The embryo had a thickened outer periclinal cell wall.
• Pericarp and aril are not essential to prevent water loss in seeds, but do help to retain seed moisture, favouring viability maintenance and promoting germination during the rainy season. Morphoanatomical features of seeds are suggested as main factors that reduce water loss. Survival of these desiccation‐sensitive seeds upon dispersal during the dry season appears to be facilitated by multiple diaspore features that prevent viability loss.

     

铁核桃根与叶水浸提液对果园间作绿肥植物的化感作用及其生理机制

以西南地区具有代表性的16种绿肥植物为受体材料,采用培养皿药膜法研究了铁核桃(Juglans sigillata)根水浸提液对受体种子发芽率及幼苗鲜重、干重的化感效应;并进一步研究了铁核桃根、叶水浸提液和胡桃醌对化感效应存在明显差异的4种绿肥植物(绿豆、红三叶、白三叶、花生)种子萌发与幼苗生长以及抗氧化酶特性的影响,以筛选适宜中国西南地区核桃园种植的绿肥植物,探讨核桃根和凋落物对绿肥作物的化感作用机制。结果表明:(1)铁核桃根水浸提液对绿豆的发芽率没有影响,但对绿豆幼苗鲜重和干重有显著抑制作用,而对其他15种绿肥的发芽率和鲜重、干重均有抑制作用。(2)胡桃醌显著抑制绿豆种子萌发,而铁核桃根或叶水浸提液对绿豆种子萌发没有影响。(3)铁核桃根或叶水浸提液以及胡桃醌对绿肥植物幼苗生长的化感效应趋势一致,但核桃根或叶水浸提液的化感效应强于胡桃醌。(4)绿豆幼苗在铁核桃根或叶水浸提液以及胡桃醌处理下,超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)的活性均高于其他3种(红三叶、白三叶、花生)受体幼苗,表明绿豆清除活性氧能力高,细胞受损害程度较低,受化感作用影响最弱。研究认为,绿豆为适宜中国西南地区幼龄核桃园种植的间作绿肥植物。     

Root growth and statocyte polarity in lentil seedling roots grown in microgravity or on a slowly rotating clinostat

The ability of clinostats to simulate microgravity was evaluated by comparing lentil ( Lens culinnrias L. cv. Verte du Puy) seedlings grown in space (Spacelab D1 Mission) with seedlings grown on a slowly rotating elinostat. Seeds were germinated and incubated for 25.5 h at 22°C (1) in microgravity, (2) on a 1g-centrifuge in space. (3) on a slowly rotating elinostat and (4) on the ground. Morphological (root length and orientation) and ultrastructural (distribution of amyloplasts, location of the nucleus in statocytes) parameters were studied. For clinostat experiments, two different configurations were employed: the longitudinal axis of the root was parallel (horizontal elinorotation) or perpendicular (vertical elinorotation) to the axis of rotation. the same configurations were used for the lg-controls. Root length and orientation were similar for roots grown on the clinostat and in microgravity. The amyloplasts were identically distributed in statocytes of horizontally clinorolated roots and in statocytes differentiated in microgravity. However, the location of the nucleus was similar in vertically rotated roots and microgravity samples. Since the involvement of the nucleus in graviperception is not known, it can be concluded that horizontal clinorotation simulates microgravity better than vertical elinorotation.     

Root nitrogen concentration of sorghum above 2% produces least Striga hermonthica seed stimulation

A series of pot and laboratory experiments was carried out to assess the effects of N status of sorghum roots and timing of N application (as NH₄NO₃) on the germination of Striga hermonthica seeds. Root N concentrations varied from 10 to 26 mg N g^?1. The cut root and the root exudates technique used in assaying S. hermonthica seed germination gave similar results. However, the cut root technique was easier to handle and more discriminating at low germination levels. S. hermonthica seed germination per unit sorghum root mass followed a broken‐stick model. It decreased with increasing root N concentrations, reaching lowest levels at a root N concentration of 19.5 mg N g^?1, after which no further reduction occurred. It was not possible to reduce S. hermonthica seed germination to a zero level. Timing of N application influenced the time a higher N concentration is reached, not the S. hermonthica seed germination. Both timing and rate of N application are important in maintaining root N concentrations above 19.5 mg N g^?1, thereby potentially reducing S. hermonthica germination in the field. Translation of results to reductions in infection levels and yield losses is hampered by density‐dependent relations after the S. hermonthica germination stage.     

Changes in plant growth processes under microgravity conditions simulated by a three-dimensional clinostat

We developed a three-dimensional (3-D) clinostat to simulate a microgravity environment and studied the changes in plant growth processes under this condition. The rate of germination of cress (Lepidium sativum), maize (Zea mays), rice (Oryza sativa), pea (Pisum sativum), or azuki bean (Vigna angularis) was not affected on the clinostat. The clinostat rotation did not influence the growth rate of their roots or shoots, except for a slight promotion of growth in azuki roots and epicotyls. On the contrary, the direction of growth of plant organs clearly changed on the 3-D clinostat. On the surface of the earth, roots grow downward while shoots upward in parallel to the gravity vector. On the 3-D clinostat, roots of cress elongated along the direction of the tip of root primordia after having changed the direction continuously. Rice roots also grew parallel to the direction of the tip of root primordia. On the other hand, roots of maize, pea, and azuki bean grew in a random fashion. The direction of growth of shoots was more controlled even on the 3-D clinostat. In a front view of embryos, shoots grew mostly along the direction of the tip of primordia. In a side view, rice coleoptiles showed an adaxial (toward the caryopsis) while coleoptiles of maize and epicotyls of pea and azuki bean an abaxial curvature. The curvature of shoots became larger with their growth. Such an autotropism may have an important role in regulation of life cycle of higher plants under a microgravity environment.     

Reactive oxygen species,ABA and nitric oxide interactions on the germination of warm-season C<Subscript>4</Subscript>-grasses

Hydrogen peroxide (H₂O₂) as a source of reactive oxygen species (ROS) significantly stimulated germination of switchgrass (Panicum virgatum L.) seeds with an optimal concentration of 20 mM at both 25 and 35°C. For non-dormant switchgrass seeds exhibiting different levels of germination, treatment with H₂O₂ resulted in rapid germination (<3 days) of all germinable seeds as compared to seeds placed on water. Exposure to 20 mM H₂O₂ elicited simultaneous growth of the root and shoot system, resulting in more uniform seedling development. Seeds of big bluestem (Andropogon gerardii Vitman) and indiangrass [Sorghastrum nutans (L.) Nash] also responded positively to H₂O₂ treatment, indicating the universality of the effect of H₂O₂ on seed germination in warm-season prairie grasses. For switchgrass seeds, abscisic acid (ABA) and the NADPH-oxidase inhibitor, diphenyleneiodonium (DPI) at 20 μM retarded germination (radicle emergence), stunted root growth and partially inhibited NADPH-oxidase activity in seeds. H₂O₂ reversed the inhibitory effects of DPI and ABA on germination and coleoptile elongation, but did not overcome DPI inhibition of root elongation. Treatment with H₂O₂ appeared to enhance endogenous production of nitric oxide, and a scavenger of nitric oxide abolished the peroxide-responsive stimulation of switchgrass seed germination. The activities and levels of several proteins changed earlier in seeds imbibed on H₂O₂ as compared to seeds maintained on water or on ABA. These data demonstrate that seed germination of warm-season grasses is significantly responsive to oxidative conditions and highlights the complex interplay between seed redox status, ABA, ROS and NO in this system.     

田菁内生菌定殖及其与多糖混合浸种的耐盐促生效果

【背景】土壤盐渍化已经成为日益严重的世界性问题,盐渍化不仅影响作物的产量,还会影响土壤的理化性质,抑制种子的萌发,阻碍植物正常生长,以及种子对水分和养分的吸收,进而影响作物的产量。【目的】玉米在盐渍土壤上生长受限,探究在中、高盐浓度下田菁种子内生菌与田菁胶混合浸种对玉米发芽的影响,为促进盐渍土玉米生长提供技术支持。【方法】利用LB液体培养基测定田菁种子内生菌贝莱斯芽孢杆菌ZH60的耐盐性;分别利用1%浓度田菁胶、OD600为0.8的ZH60菌悬液及两者混合液对玉米浸种3 h,自然风干后分别置于0、100和200 mmol/L NaCl的0.8%琼脂培养基上培养,测定玉米种子发芽势、发芽率、根长及芽长。将两叶一心期的玉米幼苗移至装有蛭石的花盆中培养,用荧光标记的内生菌ZH60灌根,分别于1、5、11、17、25 d取玉米根系研磨,利用平板菌落计数法测定内生菌在玉米根部的定殖量;利用激光共聚焦显微镜观察第28天ZH60在玉米根部的定殖情况。【结果】菌株ZH60耐11%的NaCl盐浓度,在中、高盐浓度下混合浸种的发芽势较对照组分别提高了28%、22%、30%;芽长提高了158%、163%、1...