Robin Ch, Hay M J M, Newton P C D
Institut National de la Recherche Agronomique-Ecole Nationale Supérieure d'Agronomie et des Industries Alimentaires, BP 172, F-54505, Vandoeuvre-Ies-Nancy Cedex, France.
AgResearch Grasslands, Private Bag 11008, Palmerston North, New Zealand.
Oecologia. 1994 Dec;100(3):236-242. doi: 10.1007/BF00316950.
We studied the effects of light quality and defoliation on the rate of phytomer appearance and axillary bud outgrowth in white clover. The treatments were applied to one phytomer, a phytomer being defined as the structural unit comprising a node, internode, axillary bud, subtending leaf and two nodal root primordia. Light of a low red:far-red (R:FR) ratio (0.27) was applied to a "target" phytomer either (i) within the apical bud and then to the axillary bud after emergence of the phytomer from the apical bud, or (ii) to the axillary bud only after emergence. The light conditions were directed to these specific parts of the plant by collimating light from small FR light-emitting diodes; with this technique we were able to change the light quality without any change in the level of photosynthetically active radiation. The subtending leaf of the target phytomer was retained or defoliated when it had emerged from the apical bud. FR light applied from the time the phytomer was within the apical bud caused a delay in branch appearance at the target phytomer. In contrast, direct treatment of the axillary bud with FR light after it had emerged from the apical bud did not result in any delay in branch appearance. As the light treatment of the apical bud may have changed the light environment of any of the organs contained in the bud we were unable to ascribe the delay in branch appearance to light perception by any particular organ. However, indirect evidence leads to the conclusion that the likely site of light perception was the developing leaf subtending the axillary bud while it was the outermost phytomer within the apical bud. These results do not support the hypothesis that the R:FR ratio of light incident at an axillary bud site is the environmental factor that controls bud development. Defoliation of the unfolding leaf reduced the rate of phytomer appearance on the main stolon but had no immediate effect on branch appearance. As a consequence there was a reduction in the number of phytomers between the stolon apical meristem and the first phytomer with a branch. This is frequently taken to indicate a relaxation of apical dominance, but in this case was found not to involve a direct effect on bud activity. A current model of white clover growth suggests that there is integration of activity between apical meristems but independence of activity and response to the local micro-environment by axillary buds. In contrast, we found that (i) defoliation reduced phytomer appearance only at the main stolon apical meristem and not at all the meristems in the plant and (ii) that a change in the local light environment of an axillary bud had no discernible effect on bud activity once the bud had emerged from the apical bud but could delay branching if applied before emergence. These results are at variance with the predictions of the model.
我们研究了光质和去叶处理对白三叶节间出现速率和腋芽生长的影响。处理作用于一个节间,节间被定义为包含一个节、节间、腋芽、着生叶和两个节根原基的结构单元。低红:远红(R:FR)比(0.27)的光施加于一个“目标”节间,方式为:(i)在顶芽内施加,然后在该节间从顶芽中长出后施加于腋芽;或(ii)仅在节间长出后施加于腋芽。通过准直来自小型远红光发光二极管的光,将光照条件导向植物的这些特定部位;利用该技术,我们能够改变光质,而不改变光合有效辐射水平。目标节间的着生叶从顶芽中长出后,保留或去除。从节间在顶芽内时就施加远红光,会导致目标节间的分枝出现延迟。相比之下,在腋芽从顶芽中长出后直接用远红光处理腋芽,不会导致分枝出现延迟。由于对顶芽的光照处理可能改变了芽内所含任何器官的光照环境,我们无法将分枝出现的延迟归因于任何特定器官对光的感知。然而,间接证据得出的结论是,可能的光感知部位是腋芽着生的发育中叶,而此时它是顶芽内最外层的节间。这些结果不支持以下假设:入射到腋芽部位的光的R:FR比是控制芽发育的环境因素。展开叶的去叶处理降低了主匍匐茎上节间出现的速率,但对分枝出现没有直接影响。结果,匍匐茎顶端分生组织与第一个有分枝的节间之间的节间数量减少。这通常被认为表明顶端优势的放松,但在这种情况下发现并不涉及对芽活性的直接影响。白三叶生长的当前模型表明,顶端分生组织之间存在活性整合,但腋芽的活性及其对局部微环境的响应是独立的。相比之下,我们发现:(i)去叶处理仅降低了主匍匐茎顶端分生组织处的节间出现,而不是植物中所有的分生组织;(ii)一旦腋芽从顶芽中长出,腋芽局部光照环境的变化对芽活性没有明显影响,但如果在腋芽长出之前施加,则会延迟分枝。这些结果与该模型的预测不一致。