Journal of Plant Biotechnology

The Korean Society of Plant Biotechnology (한국식물생명공학회)
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Organization and function of shoot apical meristem affecting growth and development in plants

식물의 생장과 발달에 영향을 미치는 슈트 정단분열조직의 체제와 기능

  • Lee, Kyu Bae (Department of Biological Science Education, Chosun University)
  • Received : 2014.11.14
  • Accepted : 2014.12.20
  • Published : 2014.12.31
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Abstract

In plants, a shoot apex has a small region known as the shoot apical meristem (SAM) having a group of dividing (initiating) cells. The SAM gives rise to all the groundabove structures of plants throughout their lifetime, and thus it plays important role in growth and development of plants. This review describes theories to explain the SAM organization and function developed over the last 250 years. Since in 1759 German botanist C. F. Wolff has described firstly the SAM, in 1858 Swiss botanist C. N${\ddot{a}}$geli proposed the apical cell theory from the observation of a large single apical cell in the SAM of seedless vascular plants: however, this view was recognized to be unsuitable to seed plants. In 1868, German botanist J. Hanstein suggested the histogen theory: this concept subdividing the SAM into dermatogen, periblem, and plerome was unable to generally apply to seed plants. In 1924, German botanist A. Schmidt proposed the tunica-corpus theory from the examination of angiosperm SAM in which two parts show different planes of cell division: this theory was proved to be not suitable to gymnosperm SAM, not have stable surface tunica layer. In 1938, American botanist A. Foster described zones in gymnosperm SAM based on the cytohistologic differentiation and thus called it a cytohistological zonation theory. With works by E. Gifford, in 1954, this zonation pattern was demonstrated to be also applicable to angiosperm SAM. As another theory, in 1952 French botanist R. Buvat proposed the m${\acute{e}}$rist${\grave{e}}$me d'attente (waiting meristem) theory: however, this concept was confuted because of its negation of function during vegetative growth phase to central initial cells. Rescent studies with Arabidopsis thaliana have found that formation and maintenance of the SAM are under the control of selected genes: SHOOTMERISTEMLESS (STM) gene forms the SAM, and WUSCHEL (WUS) and CLAVATA (CLV) genes function in maintaining the SAM; signaling between WUS and CLV genes act through a negative feedback loop.

식물에서 슈트 정단부에는 분열하는 세포들로 이루어진 슈트 정단분열조직이라는 조그마한 부분이 있다. 슈트 정단분열조직은 그 식물의 일생을 통해서 땅 위의 모든 구조를 만들어 낸다. 이 연구의 목적은 지난 250년 동안 슈트 정단분열조직의 체제와 기능을 설명하기 위해 발전해 온 이론들을 기술하여, 식물의 생장과 발달에 영향을 미치는 슈트 정단분열조직의 중요성을 이해하는 데 있다. 1759년 볼프(C. F. Wolff)가 최초로 슈트 정단분열조직을 기재한 이후, 1858년에 네겔리(C. N${\ddot{a}}$geli)는 무종자 유관속식물의 슈트 정단분열조직에서 하나의 커다란 정단세포를 관찰하여 정단세포설을 제안하였다. 그러나 이 설은 종자식물에 적용할 수 없는 것으로 확인되었다. 이어서 1868년 한스타인(J. Hanstein)에 의해 조직원설이 제한되었으나, 이 설도 종자식물에 일반적으로 적용할 수 없었다. 그 후, 1924년 슈미트(A. Schmidt)은 피자식물의 슈트 정단분열조직에서 세포의 분열면이 서로 다른 것을 관찰하여 초층-내체설을 제안하였다. 이 설은 나자식물에 적용할 수 없는 것으로 확인되었다. 1938년 포스터(A. Foster)는 세포조직학적으로 서로 다른 구역으로 이루어진 나자식물의 슈트 정단부를 관찰하여 세포조직학적 구역화설을 제안하였다. 또한 1954년 기포드(E. Gifford)의 연구에 힘입어 피자식물의 슈트 정단분열조직에도 세포조직학적 구역화설이 적용될 수 있다는 사실이 증명되었다. 또 다른 설로서, 1952년 뷔바(R. Buvat)는 대기 분열조직설이 제안되었으나, 영양생장 시기에 분열조직 중앙의 시원세포들이 분열하지 않는다는 주장이 수용되지 않았다. 최근에 애기장대(Arabidopsis thaliana)를 이용한 연구에서, 슈트 정단분열조직의 형성과 유지가 몇 개의 유전자들에 의해서 조절된다는 사실이 밝혀졌다. 즉, SHOOTMERISTEMLESS(STM) 유전자는 슈트 정단분열조직을 형성하고, WUSCHEL(WUS) 및 CLAVATA (CLV) 유전자들은 슈트 정단분열조직을 유지시키며, WUS와 CLV 두 유전자들 사이의 신호전달은 음성되먹임회로를 통해서 이루어진다.

Keywords

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