synthesize DNA & other molecules unless the cell is determined to enter S-phase. Its the same with cell in G2, not necessary to prepare for mitosis unless the cell is going to divide. Cells only arrested in G1 & G2 but not in M and S ( Vant Hoff, 1974). Cont. Vant Hoff also showed that cells of Pisum which were arrested in G1 can be manipulated to stop in G2. Substance/chemical originated from the cotyledon which made the cell stop/arrested in G2. ( G2 Factor- Trigonelline ( Evans et.al., 1979). Cells will stop in G2 when this trigonelline is present or stop in G1 when this substance is absent. Most legumes, Pisum sativum, Vicia faba, Glycine max and Phaseolus vulgaris contain high concentration of this substance ( Trigonelline). While non-legumes & monocots contain very low concentration of trigonelline ( Evans & Tramontano, 1984). This indicates that protein synthesis possibly involves in this mechanism. Another Cell Cycle Hypothesis Proposed by Brown ( 1976)- Genes controlling events in Cell cycle were arranged in groups. Each group correspond/related to 4 different phases (G1,G2, M, S). Only 1 gene in each group operating at any one time. Every group is controlled by gene- Gene Master Ref; Cell division in higher Plants (Yeoman,1976) Cell Cycle in Vitro Eriksson ( 1967) was the first person using cell cycle approach to study plant cells in vitro. He analysed the cell cycle in cell suspension culture of Haplopappus gracilis. Since then the accumulation of cell cycle data from plant tissues in vitro was very few. Studies on cell cultured in vitro was to understand the cell cycle system in intact/in vivo plants. In few limited studies, comparison was made between the duration of cell cycles & component phases in vivo & in vitro. Cont. Duration of Cell cyle in vitro usually always longer, mostly due to G1. Bayliss (1975) determined the duration of cell cycle in Daucus carota in the seedlings roots & also in diploid cell suspension culture supplemented with 2,4-D. He obtained the duration of cell cycle in vivo as 7.5 h and 51.2 h for cells in vitro. The duration of G1 was 39.6 h in vitro & 1.3 h in vivo. Studies on root cells of Zea mays ( Verma, 1980) showed that the duration of cell cycle was 9.9 h. Gould (1984) obtained 37 h for cells in vitro. Cont.
The duration of G1 was 23.9 in vitro
and only 1.7 h in root meristem cells. These differences portrayed differences in morphology of the meristematic cells in vivo as compared to cells in vitro. The duration of cell cycle in vivo & in vitro can be summarised as follows; Table 1
Species Duration G1 (h) S (h) G2 (h) M (h) Reference
cells Cell Cycle Regulation in Plants Cell division plays a crucial role during all phases of plant development. The molecular analysis of cell division & its regulation in plants lags far behind yeast & animals. Genes such as ras and p53 are players in yeast & animals that activate the cell-cycle engine. In all eukaryotes, the biochemical machine that controls progress through the cell cycle consists of a catalytic protein kinase & an activating cyclin subunit. This complex but neither protein alone, has protein kinase activity. Stepwise changes specifically regulate progression through the cycle. Cont. The proteins encoded by S.cerevisiae CDC28 and the S. pombe Cdc2 genes are the prototypic CDKs required at both the G1/S and G2/M transitions. In animals CDK1(=cdc2) is essential for the G2/M transition, whereas CDK2 & CDK3 are required for the G1/S transition. These 3 CDKs share a short amino acid sequence PSTAIRE Cont. Several related kinases with homology but incomplete sequence identity to the PSTAIRE domain have been identified. Cyclins were the first identified in marine invertebrates as proteins, when injected into frog oocytes could induce meiosis. The observation that S. pombe cdc13 gene, which encodes a cyclin, genetically interacts with cdc2 established a tight functional link between the components of the cell cycle engine. Since then, an increasing no. of cyclins have been identified in eukaryotes. Cont. In S. cerevisiae 3 cyclins ( CLN1, CLN2,CLN3) are expressed & presumably function in G1, 2 are expressed in S phase ( CLB5, CLB4) & 4 are expressed in mitosis ( CLB1, CLB2, CLB3, CLB4). In animals, the C-D- and E- class cyclins are expressed in G1, cyclin A is expressed in the S &G2, B cyclins are expressed in the G2 & early M phases. In most cells the decision whether to commit to division is made during G1, by G1 cyclins. Cont.
Cyclins are involved in
determining the substrate specificity of CDKs as well as in targeting CDK activity to specific subcellular compartments during the cell cycle. Cell Division in Plants Plant genes homologous to components of the cell cycle engine have been cloned recently but the function of these putative cell cycle regulators has not yet been examined in plants. Therefore, at present the functions of these genes are unknown. CDKs have been cloned from pea, alfalfa, maize, Arabidopsis. Plant cyclins have been cloned from carrots, soybean, alfalfa & Arabidopsis. The soybean & Arabidopsis gene products promote nuclear envelope breakdown when the corresponding mRNAs are injected into Xenopus oocyctes, indicating their function as mitotic cyclins. Cont. All plant cyclins share homology to both A- & B- type cyclins. It is unclear whether these plant cyclins play roles during S phase as reported for animal A type cyclins, as well as during the G2 & M phases as shown for B-type cyclins. A-type cyclins have been found only in animals. Therefore, plant AB- cyclins may be early members of an evolutionarily more divergent cyclin gene family. No G1 cyclin has yet been reported from plants. Cont.
The expression of cdc2 homologs
is higher in mitotically active tissues such as floral buds and low or undetectable in mature, nonproliferating organs such as leaves.