Wagner E, Mohr H
Botanisches Institut der Universität Freiburg i. Br., Freiburg i. Br, Deutschland.
Planta. 1966 Sep;71(3):204-21. doi: 10.1007/BF00384883.
Anthocyanin synthesis of the mustard seedling (Sinapis alba L.), a typical phytochrome-dependent photoresponse has been further investigated. - It has been found that only two types of tissue can synthesize anthocyanin under the influence of active phytochrome (=P730), namely, the epidermis of the votyledons and the subepidermal layer in the hypocotyl (Fig. 2, 3). - Under our standard conditions (25° C; cf. methods) phytochrome-potentiated anthocyanin synthesis is only possible 24 hours after sowing and it ceases about 60 hours after sowing, independent of the amount of anthocyanin which has been accumulated (Fig. 5, 6). On the basis of the whole seedling the highest sensitivity of the anthocyanin producing system to light is around 36 hours after sowing (Fig. 8). Within the tissues which are capable of forming anthocyanin there is a characteristic shift of the ability to respond to P730 as the seedling ages. If we devide the seedling into 4 segments (Fig. 9) it turns out that in the basal and middle part of the hypocotyl the ability to form anthocyanin is rapidly lost whereas in the upper part of the hypocotyl and in the cotyledons this ability even increases at first. The following decrease is slower than in the basal parts (Fig. 10, 11).It is argued that this specific and dynamic cellular pattern of responsiveness to P730 can be regarded as a manifestation of a "primary differentiation" in the course of which the genotype of each individual cell in the dark-grownt seedling is devided into 3 functional types of genes: active, inactive, and potentially active genes (P730) (Fig. 4). - In connection with anthocyanin synthesis P730 is thought to act exclusively at the level of "secondary differentiation", i.e., it is thought to initiate the action of potentially active genes via a signal-chain. The action of P730 is non-specific. The specificity of the photoresponse of an individual cell is determined by the status of its "primary" differentiation (Fig. 4).If the process of differentiation is slowed down (e.g. by the application of low doses of Actiomycin D) anthocaynin synthesis can continue much longer than under our standard conditions where it ceases around 60 hours after sowing (Fig. 12). This fact seems to indicate that the loss of the ability to form anthocyanin is due to an inactivation of pertinent genes by the process of "primary differentiation", which is itself, as one would expect, under the control of genes.
对典型的依赖光敏色素的光反应——芥菜幼苗(白芥)的花青素合成进行了进一步研究。已发现,在活性光敏色素(=P730)的影响下,只有两种组织能够合成花青素,即子叶表皮和下胚轴的亚表皮层(图2、3)。在我们的标准条件下(25℃;参见方法),光敏色素增强的花青素合成只有在播种后24小时才可能发生,并且在播种后约60小时停止,与积累的花青素量无关(图5、6)。基于整株幼苗,花青素产生系统对光的最高敏感性在播种后约36小时左右(图8)。在能够形成花青素的组织内,随着幼苗年龄的增长,对P730作出反应的能力存在特征性变化。如果我们将幼苗分成4段(图9),结果表明,在下胚轴的基部和中部,形成花青素的能力迅速丧失,而在下胚轴上部和子叶中,这种能力起初甚至会增加。随后的下降比基部慢(图10、11)。有人认为,这种对P730的特异性和动态细胞反应模式可被视为“初级分化”过程的一种表现,在此过程中,黑暗中生长的幼苗中每个细胞的基因型被分为3种功能类型的基因:活性基因、非活性基因和潜在活性基因(P730)(图4)。与花青素合成相关,P730被认为仅在“次级分化”水平起作用,即它被认为通过信号链启动潜在活性基因的作用。P730的作用是非特异性的。单个细胞光反应的特异性由其“初级”分化状态决定(图4)。如果分化过程减慢(例如通过施用低剂量放线菌素D),花青素合成可以比我们的标准条件下持续长得多的时间,在标准条件下,它在播种后约60小时停止(图12)。这一事实似乎表明,形成花青素能力的丧失是由于“初级分化”过程中相关基因的失活,而“初级分化”过程本身正如人们所预期的那样,受基因控制。