Auxin

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N. Shinohara (shino@nibb.ac.jp) , Y Hiwatashi, T. Aoyama, N. Aono, T. Hirai, and T. Nishiyama (tomoakin@kenroku.kanazawa-u.ac.jp)


Contents

Auxin biosynthesis and metabolism

Auxin is involved in various aspects of plant development (Murphy 2002). CYP79B2, CYP79B3, CYP83B1/SUR2, NIT1, NIT2, NIT3, YUCCA, and AAO operate in the Trp-dependent biosynthesis pathways (Woodward and Bartel, 2005a). Potential orthologs except the last two genes were not found in S. moellendorffii and P. patens. This suggests the diversity of Trp-dependent pathways among land plant lineages. Endogenous IAA levels are also controlled by metabolisms of IAA conjugates. Seven of 19 members in A. thaliana GH3 gene family are responsible for IAA-amide formation (Staswick et al., 2005). Potential orthologs of DFL1 of this family were found in all the land plant lineages and angiosperms have more potential orthologs than S. moellendorffii and P. patens have. This is concordant with the lower levels of IAA conjugates in basal land plants than angiosperms (Sztein et al., 1995).


Auxin signaling

The wide array of auxin actions in flowering plants is mediated by three groups of proteins: TIR1/AFB F-box proteins that perceive auxin molecules, Aux/IAA transcription repressors, and ARF transcription activators (Weijers and Jurgens, 2004). We found all the three gene families in each land plant, suggesting the conservation of auxin signaling system in land plants. TIR1 functions as an auxin receptor and COI1 presumably functions as a jasmonic acid receptor (Woodward and Bartel, 2005b). Phylogenetic trees of their homologs showed that TIR1 and COI1 orthologs diverged before the divergence of P. patens from other land plants. This is concordant with the divergence time of two GH3 gene groups, the DFL homologs that function in jasmonic acid and IAA-amido synthases. The number of Aux/IAA genes was conspicuously increased in the angiosperm lineage after the branching of S. moellendorffii, and we found four and two Aux/IAA potential orthologs in S. moellendorffii and P. patens (Table). The change in the number of Aux/IAA genes likely influenced the auxin signaling systems in land plants. Twenty-three members of A. thaliana ARF genes were classified into three groups (Okushima et al., 2005), and S. moellendorffii and P. patens have potential orthologs for all the groups. The numbers of ARF homologs were increased in each angiosperm, S. moellendorffii, and P. patens lineage, and as a result, each group has similar number of ARF genes in each genome.

Auxin carriers

An auxin polar transport is necessary for a proper development in all land plants examined (Cooke et al., 2002). AUX/LAX genes are involved in auxin influx to a cell, and PIN and PGP/MDR genes are in efflux. These carriers are necessary for a proper polar auxin transport. We could find AUX/LAX, PGP/MDR, and PIN potential orthologs in S. moellendorffii and P. patens. A. thaliana PIN paralogs function in different organs. PIN5 phylogenetic tree showed that they diverged in the angiosperm lineage after the branching of S. moellendorffii, although the bootstrap supports are not high. AUX paralogs were also diverged in each angiosperm, S. moellendorffii, and P. patens lineage.


References

Murphy A (2002) Auxin: the growth hormone. In L Taiz, E Zeiger, eds, Plant Physiology, Ed 3. Sinauer Associates Inc. Publishers, Sunderland, MA, pp pp 423–460

Cooke, T.J., Poli, D., Sztein, A.E., and Cohen, J.D. (2002). Evolutionary patterns in auxin action. Plant Mol Biol 49, 319-338.

Okushima, Y., Overvoorde, P.J., Arima, K., Alonso, J.M., Chan, A., Chang, C., Ecker, J.R., Hughes, B., Lui, A., Nguyen, D., Onodera, C., Quach, H., Smith, A., Yu, G., and Theologis, A. (2005). Functional genomic analysis of the AUXIN RESPONSE FACTOR gene family members in Arabidopsis thaliana: unique and overlapping functions of ARF7 and ARF19. Plant Cell 17, 444-463.

Staswick, P.E., Serban, B., Rowe, M., Tiryaki, I., Maldonado, M.T., Maldonado, M.C., and Suza, W. (2005). Characterization of an Arabidopsis enzyme family that conjugates amino acids to indole-3-acetic acid. Plant Cell 17, 616-627.

Sztein, A.E., Cohen, J.D., Slovin, J.P., and Cooke, T.J. (1995). Auxin metabolism in representative land plants. American Journal of Botany 82, 1514-1521.

Weijers, D., and Jurgens, G. (2004). Funneling auxin action: specificity in signal transduction. Curr Opin Plant Biol 7, 687-693.

Woodward, A.W., and Bartel, B. (2005a). A receptor for auxin. Plant Cell 17, 2425-2429.

Woodward, A.W., and Bartel, B. (2005b). Auxin: regulation, action, and interaction. Ann Bot (Lond) 95, 707-735.


Table of gene numbers in each species

Gene functions Gene Gene used as a query The number of putative orthologs
Arabidopsis thaliana Oryza sativa Selaginalla moellendorffii Physcomitrella patens
Auxin metabolism YUCCA YUCCA 9 8 3 (6) 6
Auxin metabolism CYP79B2 and B3 1) CYP79B2 2 0 0 0
Auxin metabolism NIT1-3 NIT1 3 0 0 0
Auxin metabolism CYP83B1/SUR2 1) SUR2 2 0 0 0
Auxin metabolism GH3.6/DFL1 2) GH3.6/DFL1 19 11 17 (29) 2
Auxin metabolism AAO1 AAO1 4 6 3 (5) 2
Auxin metabolism TYDC1, 2 TYDC2 2 1 5 (9) 2
Auxin metabolism CYP79F1/SPS1 SUPERSHOOT1 8 3 0 0
Auxin signalling TIR1 TIR1 6 6 2 (4) 4
Auxin signalling ABP1 ABP1 1 1 2 (4) 1
Auxin signalling IAA1 to 20, 26 to 34 2) IAA31 29 30 4 (7) 2
Auxin signalling ARF5/MP, NPH4/BIPOSTO, ARF6, 8, 16, and 19 2) NPH4 5 8 3 (5) 6
Auxin signalling ETT/ARF3, ARF1, 2, 4, 9, 11 to 15, 18, 20, and 21 ETT 15 9 2 (4) 4
Auxin signalling ARF10, ARF16 ARF16 2 4 2 (4) 2
Auxin signalling ARF17 ARF16 1 1 0 0
Auxin signalling AXR1 2) AXR1 3 1 1 (2) 2
Auxin signalling SKP1/ASK1 SKP1 17 12 1 (2) 3
Auxin signalling AXR6/ATCUL1 ATCUL1 2 3 1 (2) 3
Auxin signalling RCE1 RCE1 2 3 3 (5) 5
Auxin carriers AUX1 AUX1 4 4 2 (4) 4
Auxin carriers PIN1 to 8 2) PIN5 9 11 5 (7) 3
Auxin carriers MDR/PGP1 PGP1 1 1 0 0
Auxin carriers PGP19 PGP1 1 2 2 (4) 2
Auxin carriers MDR4/PGP4 PGP1 8 8 5 (9) 1
Auxin carriers PID PID 3 3 0 0
Auxin carriers BIG1 BIG1 1 1 1 (2) 2

Table of gene models in the assembly

protein ID name named by other name by location comm
23133 YUC1-1
64527 YUC1-2
102211 YUC2-1 YUC2 Naoki Shinohara
80431 YUC2-2
75206 YUC3-1
87220 YUC3-2
99818 DFL1-1
103740 DFL1-2
125904 DFL1L1-1
129356 DFL1L1-2
235392 DFL1L2-1
131591 DFL1L2-2
74871 DFL1L3-1
87623 DFL1L3-2
163668 DFL1L4A-1 Selmo1/scaffold_0:1498358-1500323
268799 DFL1L4A-2 Selmo1/scaffold_60:98295-100261
73348 DFL1L4B-1 Selmo1/scaffold_0:1492572-1494472
119116 DFL1L4B-2 Selmo1/scaffold_60:322081-323988
110439 JAR1-1
131513 JAR1-2
234778 JAR1L1-1
125443 JAR1L1-2
416246 JAR1L2-1
235186 JAR1L2-2
31681 JAR1L3-1
31683 JAR1L3-2
77060 JAR1L4-1
138788 JAR1L4-2
437924 JAR1L5-1
441348 JAR1L5-2
115497 JAR1L6-1
21596 JAR1L6-2
125145 JAR1L7-1 Selmo1/scaffold_77:894911-896539 left
128888 JAR1L7-2 Selmo1/scaffold_92:82755-84383 left
124989 JAR1L8-1 Selmo1/scaffold_77:885872-887788 right
427552 JAR1L8-2 Selmo1/scaffold_92:74324-76231 right
89728 TYDC1-1 Selmo1/scaffold_10:416396-418439
171450 TYDC1-2 Selmo1/scaffold_10:425812-427870
89475 TYDC2-1 Selmo1/scaffold_10:425812-427870
121532 TYDC3-1
123605 TYDC3-2
79134 TYDC4B Selmo1/scaffold_2:1849208-1851382
78872 TYDC4A Selmo1/scaffold_2:559748-561523 suspected allelic variants, although on the same scaffold
84876 TYDC5-1 Selmo1/scaffold_6:2373505-2375717
169665 TYDC5-2 Selmo1/scaffold_10:389236-391616
109785 ABP1-1 Naoki Shinohara right
133599 ABP1-2 Selmo1/scaffold_120:47761-48411
451561 ABP2-1 Selmo1/scaffold_39:626483-627248
451562 ABP2-2 Selmo1/scaffold_120:48614-49385
166729 ETT1-1 Selmo1/scaffold_4:1852401-1856602
430287 ETT1-2 Selmo1/scaffold_127:187674-190881
437944 ETT2-1 Selmo1/scaffold_1:4121931-4125437
438973 ETT2-2 Selmo1/scaffold_5:122023-123827
437622 ARF10A-1
451564 ARF10A-2
443304 ARF10B-1
444549 ARF10B-2
103350 AXR1-1
126167 AXR1-2
271223 SKP1-1
154042 SKP1-2
173394 CUL1-1
158171 CUL1-2
98006 CUL3-1
136955 CUL3-2
450664 RCE1-1 RCE2-4 Ying Li
450663 RCE1-2 RCE2-3 Ying Li
450656 RCE2-1 Ying Li RCE2-1 Ying Li
450657 RCE2-2 Ying Li RCE2-2 Ying Li
121306 RCE3-2
141341 BIG1-1
153843 BIG1-2
439204 AUX1A-1
176094 AUX1A-2
165624 AUX1B-1
112710 AUX1B-2
170974 TIR1A-1
178850 TIR1A-2
168175 TIR1B-1
179436 TIR1B-2
234325 PIN1-1
131936 PIN1-2
102666 PIN2-1
443592 PIN2-2
98910 PIN3-1
105586 PIN3-2
422990 PIN4-1
129093 PIN4-2
88887 PIN5-1
451571 PIN5-2
410515 PGP19A-1 SmABCB4 Uener Kolukisaoglu Selmo1/scaffold_13:77332-81556
92485 PGP19B-1 SmABCB5 Uener Kolukisaoglu Selmo1/scaffold_13:82091-86435
138662 PGP19A-2 Selmo1/scaffold_301:7964-12402
114581 PGP19B-2
169182 PGP4A-1 SmABCB7
154740 PGP4A-2
123915 PGP4B-1 SmABCB8
184079 PGP4B-2
123936 PGP4C-1 SmABCB9
184091 PGP4C-2
74892 PGP4D-1 SmABCB10
87743 PGP4D-2
176522 PGP4E-1 SmABCB6
177681 PGP4E-2
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