Experimental investigations of the shoot apex of Dryopteris aristata Druce

Abstract

A method whereby the apical meristem of the fern Dryopteris aristata Druce can be partially isolated from the adjacent lateral organs and tissues is described. This procedure has been adopted as a means of investigating growth and morphogenesis at the shoot apex. The technique involves the severance of the incipient vascular tissue which originates immediately below the apical meristem; the isolated meristem is thus seated on a plug of growing medullary parenchyma. Leaf primordia can be similarly isolated. Meristems treated in this way are capable of growth. They develop into short vasculated shoots bearing leaves. The nutrients sustaining this growth must reach the apical meristem from below by diffusing through medullary parenchyma at the base of the isolated terminal region. Above the parenchymatous region a solenostelic vascular system is present in the new axis; this is in marked contrast to the dictyostelic configuration of the parental shoot below. On the further growth of the isolated meristem leaves are produced and the stele becomes dictyostelic. The new leaves, of which as many as fourteen have been observed after 11 weeks’ growth, show the normal phyllotactic arrangement, and this is continuous with that of the main shoot below. The procedure adopted has the effect of removing the physiological dominance of the apical meristem relative to the main shoot; thus numerous large buds develop on the lateral segments of the parental shoot but none on the isolated terminal region. The growth of isolated leaf primordia is very limited. The vascular system develops as a solenostele, foliar gaps are not formed in the region of confluence with the shoot stele, axillary buds are developed, and the leaf apex becomes directed outwards. These several features are in marked contrast to the normal development. The isolated lateral segments are also capable of further growth. The experimental procedure adopted involves the severance of the vascular tissues at various levels. An account is given of new and hitherto unrecorded morphological developments observed in these segments. Interesting features include the formation of large solenostelic buds, the solenostelic development of isolated meristeles, medullation of meristeles and the induction of a polycyclic stelar condition, in one instance by a process of cambium-like activity. These are all in marked contrast to the normal development of the intact shoot. The data which have been obtained are discussed with special reference to the path of translocation of nutrients to the terminal meristem and to leaf primordia, morphogenetic processes at the shoot apex, the factors influencing the differentiation of the vascular system, and theories of shoot formation and constitution. The results of these experiments give no support to phytonic theories but emphasize the difference in potentiality for development between shoot and leaf primordia. In this connexion the factors which determine the shape and system of segmentation of the apical initials of shoot and leaf are seen to require further investigation. The hypotheses that lateral buds are inhibited by substances proceeding from the apical meristem, that the initial differentiation of vascular tissue can be attributed to the basipetal diffusion of a substance or substances from the actively growing apical meristem, and that under conditions of tensile stress incipient vascular tissue undergoes a parenchymatous development, are supported by the data of these experiments. The observations afford a clear indication of the diversity of the morphogenetic activity in the growing region. Nutritional, mechanical and other factors are seen to be important in influencing the distribution of tissues during development. The view entertained by comparative morphologists that the vascular system in ferns is of a highly conservative nature and therefore of great value in phyletic studies is to some extent opposed by the data of these experiments. But notwithstanding the several unusual vascular configurations produced as a result of the experimental treatment, there is eventually a return to the typical vascular arrangements of the normal shoot. There is thus a need for harmonizing the data of the causal and phyletic aspects. The more thoroughly the operation of morphogenetic factors extrinsic to the specific hereditary substance is understood, the more critical will be the selection of criteria of comparison for phyletic purposes.