Meristems

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Contents

Shoot meristem

Y. Hiwatashi, T. Aoyama, M. Iwata, S. Miyazaki, S.-I. Morinaga, N. Onodera, N. Sumikawa, and T. Nishiyama

hiwatash@nibb.ac.jp, tomoakin@kenroku.kanazawa-u.ac.jp

An indeterminate shoot meristem in the diploid generation is a vascular plant-specific developmental characteristic (Raven et al., 2005); however, P. patens forms an indeterminate shoot meristem in its haploid generation. In A. thaliana, the indeterminate meristem at the shoot apex is maintained by a feedback loop composed of WUS and the CLV1, 2 and 3 genes (Williams and Fletcher, 2005). We found possible WUS land plant orthologues in both S. moellendorffii and P. patens, whilst CLV1 and 2 orthologues were found in S. moellendorffii but not in P. patens. These suggest that a CLV-WUS feedback loop does not function in the indeterminate meristems of P. patens, although several genes paralogous to the CLVs may serve those functions. In all of the land plant lineages, we found transcription factors and chromatin factors that function in WUS regulation: PHB, PHV, REV, and HAN as the former and NFB2/FAS1, NFB1/MUB3/FAS2, and CHR3/SYD as the latter. It remains to be determined whether these genes regulate WUS homologue transcription in S. moellendorffii and P. patens. The class 1 KNOX genes encode essential transcription factors for the indeterminate shoot apical meristem (Williams and Fletcher, 2005), and their regulation by AS1 and AS2 is indispensable for proper leaf development. The class 1 KNOX gene expression is partly regulated by CUC transcription factors. We found putative class 1 KNOX genes and CUC orthologues in all of the land plant lineages, whilst putative AS1 orthologues were found in S. moellendorffii but not in P. patens. No putative AS2 orthologues were found in S. moellendorffii nor P. patens. These data suggest that the genes regulating shoot apical meristem development vary in land plants.


Axillary meristem

Y. Hiwatashi, T, Aoyama, and T, Nishiyama

hiwatash@nibb.ac.jp, tomoakin@kenroku.kanazawa-u.ac.jp

The emergence of a branched sporophyte is one of the key evolutionary events in land plants (Kenrick and Crane, 1997). Bryophyte sporophytes do not branch, whilst extant vascular plants have branched sporophytes, although lycopods have a bifurcated branch system, which is different from the seed plant branch system with axillary meristems. Putative orthologues of the A. thaliana MAX1, 2, 3, and 4 genes, which regulate axillary branching (Bennett et al. 2006) were found in S. moellendorffii, whilst all except MAX1 were present in P. patens. The RAX and LAS genes encode transcription factors that function in axillary meristem development (Muller et al., 2006). We found putative orthologues of LAS in both S. moellendorffii and P. patens and putative orthologues of RAX in S. moellendorffii. These results indicate that most genes for axillary meristem development existed before the evolution of the axillary meristem, and that the evolution of their interaction might have been important for the development of the new trait.

References

Bennett T, Sieberer T, Willett B, Booker J, Luschnig C, Leyser O. The Arabidopsis MAX pathway controls shoot branching by regulating auxin transport. Curr Biol. 2006 Mar 21;16(6):553-63.

Kenrick, P., and Crane, P.R. (1997). The Origin and Early Diversification of Land Plants: A Cladistic Study. (Washington, DC: Smithsonian Institution Press).

Muller, D., Schmitz, G., and Theres, K. (2006). Blind homologous R2R3 Myb genes control the pattern of lateral meristem initiation in Arabidopsis. Plant Cell 18, 586-597.

Raven, P.H., Evert, R.F., and Eichhorn, S.E. (2005). Biology of Plants. (New York: W. H. Freeman and Company).

Williams, L., and Fletcher, J.C. (2005). Stem cell regulation in the Arabidopsis shoot apical meristem. Curr. Opin. Plant Biol. 8, 582-586.



Table of gene numbers

Gene functions Gene Gene used as a query The number of putative orthologs
Arabidopsis thaliana Oryza sativa Selaginalla moellendorffii (a) Physcomitrella patens
Shoot meristem WUS and WOX1 to 14 WUS 16 10 9 (16) 3
Shoot meristem CLV1 CLV1 4 6 3 (3) 0
Shoot meristem CLV2 CLV2 2 3 2 (2) 0
Shoot meristem STM, BP, KNAT1, and 6 KNAT1 4 9 3 (3) 3
Shoot meristem KNAT3 to 5, and 7 KNAT3 4 4 2 (2) 2
Shoot meristem AS1 AS1 1 1 1 (2) 0
Shoot meristem AS2 AS2 1 2 0 0
Shoot meristem BOP1 and 2 BOP1 2 2 2 (3) 3
Shoot meristem CUC1, 2, and 3 CUC1 11 12 3 (6) 8
Shoot meristem/Gynoecium BEL1, BLR, PNF, and PNY BLR 13 13 2 (2) 4
Shoot meristem TPL TPL 5 2 3 (6) 2
Shoot meristem HAN HAN 3 2 2 (3) 4
Shoot meristem ULT1 ULT1 2 1 0 0
Shoot meristem BBM1 ANT 8 11 1 (2) 3
Axillary meristem MAX1 MAX1 1 5 1 (2) 0
Axillary meristem MAX2 MAX2 1 1 1 (2) 1
Axillary meristem MAX3 MAX3 1 1 1 (2) 3
Axillary meristem MAX4 MAX4 1 2 1 (2) 1
Axillary meristem RAX1 RAX1 6 8 1 (2) 0
Axillary meristem LAS LAS 1 2 1 (2) 2

footnote: The number of putative orthologs here refers to number of genes that is included in a clade that corresponds to all genes derived from a single gene in the last common ancestor of P. patens, S. moellendrffii, A. thaliana, and O. sativa based on phylogenetic analyses. The alignments and trees are available through http://moss.nibb.ac.jp/treedb/ (a) number of putative loci at first and number of putative alleles detected in parentheses. That is, 1 (2), indicates we found two sequences that likely represent two alleles of one locus.

Table of gene models in the assembly

protein id gene name annotated by
WUSCHEL like homeobox genes
417553 WOX5-1
419270 WOX5-2
451352 WOX2-1
404134 WOX3-1
407131 WOX3-2
451015 WOX4-1
84885 WOX1-1
451353 WOX1-2
115439 WOX6-1
438787 WOX7-1
451355 WOX7-2
404178 WOX8-1
407157 WOX8-2
451357 WOX9-1
451358 WOX9-2
KNOTTED1-like homeobox genes
415291 SmKN1-1
268085 SmKN1-2
159366 SmKN2-1
448401 SmKN2-2
90744 SmKN3-1
173857 SmKN3-2
135843 SmKN4-1
137260 SmKN4-2
CLAVATA1-related genes
146738 CLV1A-1
121783 CLV1A-2 Saori Miyazaki
186143 CLV1B-1
172284 CLV1B-2 Saori Miyazaki
83051 CLV1C-1
77558 CLV1C-2 Saori Miyazaki
CLAVATA2 like genes
425288 CLV2L1-1
128564 CLV2L1-2
BLADE ON PETIOLES related
107813 BOP1-1
125007 BOP1-2
74877 BOP2-1
91240 BOP2-2
NAM/CUC related genes
104940 NAM1-1
125472 NAM1-2
71303 NAM2-1
67245 NAM2-2
77371 NAM3-1
95019 NAM3-2
BELL-like homeobox genes
80869 BLH1-1
414952 BLH1-2
68227 BLH2-1
48157 BLH2-2
TPL related genes
404377 TPLa-1 Yuji HIWATASHI
79194 TPLa-2 Yuji HIWATASHI
163891 TPLb-1 Yuji HIWATASHI
439915 TPLb-2 Yuji HIWATASHI
88677 TPLc-1 Yuji HIWATASHI
115161 TPLc-2
HAN related genes
405209 HANa-1 Yuji HIWATASHI
415035 HANa-2 Yuji HIWATASHI
451362 HANb-1
107936 HANb-2
ASYMMETRIC LEAVES 1 / ROUGH SHEETH 2/ PHANTASTICA related genes
140962 ARP-1
270481 ARP-2
OBERON related genes
158800 OBE1-1
162596 OBE1-2
ANT related
1154 ANT-1
107288 ANT-2
MAX1 related
96541 MAX1-1 Tsuyoshi Aoyama
97512 MAX1-2 Tsuyoshi Aoyama
MAX2 related
90091 MAX2-1 Tsuyoshi Aoyama
130722 MAX2-2 Tsuyoshi Aoyama
MAX3 related
11120 MAX3-1 Tsuyoshi Aoyama
42070 MAX3-2 Tsuyoshi Aoyama
MAX4 related
173016 MAX4-1 Tsuyoshi Aoyama
127411 MAX4-2 Tsuyoshi Aoyama
RAX1 related
39509 RAX1-1 Yuji HIWATASHI
39504 RAX1-2 Yuji HIWATASHI
LAS related
84560 LAS-1
95651 LAS-2
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