Shoot apical meristem (SAM) of plants harbors stem cells capable of generating the aerial tissues including reproductive organs. Therefore, it is very important for plants to control SAM proliferation and its density as a survival strategy. The SAM is regulated by the dynamics of a specific gene network, such as the WUS-CLV interaction of A. thaliana. By using a mathematical model, we previously proposed six possible SAM patterns in terms of the manner and frequency of stem cell proliferation. Two of these SAM patterns are predicted to generate either dichotomous or axillary shoot branch. Dichotomous shoot branches caused by this mechanism are characteristic of the earliest vascular plants, such as Cooksonia and Rhynia, but are observed in only a small minority of plant species of the present day. On the other hand, axillary branches are observed in the majority of plant species and are induced by a different dynamics of the feedback regulation between auxin and the asymmetric distribution of PIN auxin efflux carriers. During evolution, some plants may have adopted this auxin-PIN system to more strictly control SAM proliferation.
SAM Pattern is Limited by Developmental ConstraintsShoot apical meristem (SAM) of plants is located at the tip of the shoot and has stem cells that generate the aerial parts including reproductive organs. Thus, it is very important for plants to control SAM proliferation and keep an appropriate density of the SAM as a survival strategy. Presumably, the fitness function would have its maximum at an optimum density
Strategy for shoot meristem proliferation in plantsHironori Fujita* and Masayoshi Kawaguchi Division of Symbiotic Systems; National Institute for Basic Biology; National Institute for Natural Sciences; Okazaki, Japan of the SAM, while the density higher or lower than this optimum would reduce the fitness (Fig. 1). Therefore, plants will evolve their SAM density toward the optimum. On the other hand, the SAM proliferation is limited by developmental constraints. The SAM development is regulated by the dynamics of a specific gene network, such as the WUSCEL (WUS)-CLAVATA (CLV) interaction of A. thaliana.1,2 We recently proposed a mathematical model based on this molecular dynamics and provided a theoretical explanation for various SAM patterns observed in plants. 3 In our model, SAM pattern is determined by two factors: mode of stem cell proliferation and stem cell containment (Fig. 2). The modes of stem cell proliferation can be classified into four groups based on regulatory strengths in the WUS-CLV dynamics: elongation mode, division mode, emergence mode and fluctuation mode (Fig. 2).3 On the other hand, the stem cell containment is associated with the spatial restriction of the WUS-CLV gene network dynamics. The combination of these two factors predicts six possible SAM patterns in terms of the manner and speed of stem cell proliferation (Fig. 2).Although our model is originally based on experimental results in A. thaliana, the outcome of the model is applicable to all plant ...