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First published online July 9, 2004; 10.1104/pp.104.041459 Plant Physiology 135:1514-1525 (2004) © 2004 American Society of Plant Biologists The ANTHER INDEHISCENCE1 Gene Encoding a Single MYB Domain Protein Is Involved in Anther Development in Rice1CSIRO Plant Industry, Canberra, Australian Capital Territories 2601, Australia; and New South Wales Agricultural Genomics Centre, Wagga Wagga, New South Wales 2678, Australia
Using a two-element iAc/Ds transposon-tagging system, we identified a rice (Oryza sativa L. cv Nipponbare) recessive mutant, anther indehiscence1 (aid1), showing partial to complete spikelet sterility. Spikelets of the aid1 mutant could be classified into three types based on the viability of pollen grains and the extent of anther dehiscence. Type 1 spikelets (approximately 25%) were sterile due to a failure in accumulation of starch in pollen grains. Type 2 spikelets (approximately 55%) had viable pollen grains, but anthers failed to dehisce and/or synchronize with anthesis due to failure in septum degradation and stomium breakage, resulting in sterility. Type 3 spikelets (approximately 20%) had normal fertility. In addition, aid1 mutant plants had fewer tillers and flowered 10 to 15 d later than the wild type. The Ds insertion responsible for the aid1 mutation was mapped within the coding region of the AID1 gene on chromosome 6, which is predicted to encode a novel protein of 426 amino acids with a single MYB domain. The MYB domain of AID1 is closely related to that of the telomere-binding proteins of human, mouse, and Arabidopsis, and of single MYB domain transcriptional regulators in plants such as PcMYB1 and ZmIBP1. AID1 was expressed in both the leaves and panicles of wild-type plants, but not in mutant plants.
Anther dehiscence is the final step of anther development and results in the release of pollen grains to enable pollination, fertilization, and seed set (Goldberg et al., 1993  ). In anthers, three tissues—endothecium, septum, and stomium—play important roles in anther dehiscence. The endothecium is an anther wall tissue lying between the epidermal layer and the tapetum. The septum is a filling tissue for spaces between vascular bundles and the two adjacent anther locules. The stomium is a single layer of specialized epidermal cells that joins adjacent anther walls and is the final breakage site for anther dehiscence (Keijzer, 1987). In rice (Oryza sativa), at the beginning of anthesis, filaments begin to elongate and pollen grains swell rapidly. The increased pollen pressure causes the locule to bulge, resulting in rupture of the septum that has been weakened by hydrolytic enzyme action. Subsequent lysis of the middle lamellae of the stomium cells, pollen pressure, and the force induced by the inward bending of the locules causes the stomium to split (Keijzer et al., 1996; Matsui et al., 1999). Finally, shrinkage of the U-shaped, thickened endothecium induces the outward bending of locules, which widens the theca to release swollen pollen grains. Meanwhile, for successful anther dehiscence and pollination, a series of developmental events that leads to the breakage of the stomium and release of pollen grains must synchronize with other developmental processes that occur within both the anther and the floret. However, little is know about the mechanisms that control and coordinate many of these developmental events. Identification of mutants defective in any of these developmental activities would shed light on the mechanism of anther dehiscence and anthesis.
Although a number of spontaneous, as well as induced, cytoplasmic and genic male sterile mutants, which have defects at different developmental stages, have been identified, few have been studied in detail at a molecular or genetic level in rice (Kinoshita, 1997 Here we report the identification and characterization of a Ds-tagged rice mutant, designated anther indehiscence1 (aid1), having defects in the development of pollen grains, endothecium strength, septum degradation, and/or stomium breakage that result in the failure of anther dehiscence and its synchronization with anthesis, leading to partial to complete spikelet sterility. Other associated phenotypes were reduced tiller numbers and delayed flowering. AID1 encodes a single MYB DNA-binding domain protein. The MYB domain of AID1 is closely related to that of telomere-binding proteins, including TRF1 and TRF2 of human and mouse, TRP1 and TBP1 of Arabidopsis, and of single MYB-repeat transcriptional regulators in plants such as MYB1 of Petroselinum crispum and IBP1 of maize (Zea mays).
Identification of Mutants with Partial to Complete Spikelet Sterile Phenotype
The initial iAc/Ds mutagenesis protocol is described elsewhere (Upadhyaya et al., 2002
Characterization of the Spikelet Sterility Mutant In the wild type, anthers reach the top of the spikelet and begin to dehisce, resulting in release of pollen grains over the stigma of the pistil immediately before spikelet opening (Fig. 1A). After anthesis, the filaments elongate further and the remaining pollen grains are released (Fig. 1B). The spikelet then begins to close, keeping empty anthers outside the spikelet (Fig. 1C). In the mutant, the anthesis of spikelets showed varying levels of abnormalities, resulting in varying levels of spikelet fertility. The predominant mutant phenotype (type 2 spikelet, see below) is shown in Figure 1, E to H. Anthers of the mutant did not dehisce at the time of spikelet opening or even after its closing (Fig. 1, E–G), while other reproductive organs appeared normal. This suggests that the spikelet sterility of the mutant was due mainly to the inability of anthers to dehisce and release pollen grains normally (compare Fig. 1, D and H). Therefore, we have designated this mutant aid1.
To better understand the variation in spikelet sterility, spikelets of the aid1 mutant were categorized into three types, based on the behavior at anther dehiscence and pollen viability. Type 1 spikelets (approximately 25%) did not show filament elongation, spikelet opening, or anther dehiscence because of abortion of pollen grains; consequently, anthers of type 1 spikelets were smaller and contained no pollen grains or only nonviable pollen grains (no or less starch). Type 2 spikelets (approximately 55%) were normal in filament elongation and spikelet opening, but showed indehiscence (Fig. 1, E–G) or delayed dehiscence, which caused spikelet sterility. There was no significant difference in the percentage of viable pollen grains between type 2 and wild-type spikelets (Table II), but pollen grains of the mutant were slightly smaller (data not shown). Furthermore, pollen grains from manually dehisced type 2 spikelets were able to self-fertilize (data not shown). Type 3 (approximately 20%) spikelets showed normal anther dehiscence and were fertile. The percentage of sterile spikelets (approximately 80%), based on this observation, was consistent with the average percentage of spikelet sterility of the homozygous aid1 mutant plants (73%–92%) shown in Table I.
No distinguishable microscopic difference could be observed between cross-sections of anthers of type 3 spikelets of the aid1 mutant and those of wild type in different stages of anther development (compare Fig. 2, B and F). Even for anthers of type 1 and 2 spikelets, no difference in the development of anthers was detected until the nonvacuolate microspore stage (data not shown). In the wild type, at the vacuolate microspore stage, the tapetal cells and the septum began to degenerate, and a cavity was formed due to the stomium dissociation from the enzymatically weakened septum (Fig. 2A). In contrast, at the same developmental stage, breakage of the septum and the formation of the cavity were not observed in anthers of type 2 spikelets (compare Fig. 2, A and D) even after complete maturation of pollen grains (compare Fig. 2, B and E). In anthers of type 1 spikelets, not only were the breakage of the septum and the splitting of the stomium blocked, but the development of pollen grains was also affected. It seemed that the development of pollen grains was arrested at different stages between the bicellular vacuolated stage and the final starch-filling stage. In most anthers, the irregularly shaped nonviable pollen grains were due to incomplete starch accumulation (Fig. 2G); in some anthers, pollen grains failed to accumulate any starch (Fig. 2H), and, in extreme cases, pollen grains degraded completely (Fig. 2I).
To determine whether environmental factors contributed to the variation of anther dehiscence and spikelet fertility, the effect of light and humidity was investigated. For dark treatment, panicles of the aid1 mutant were wrapped in black paper bags to achieve a totally dark microenvironment for the panicle. For humidity treatment, panicles of the aid1 mutant were wrapped in a plastic bag and water was injected into the bag daily with a syringe to maintain saturated humidity. Observations were made for anther dehiscence and spikelet sterility as described above. No changes in the percentage of the three types of spikelets were observed (data not shown), suggesting that light and humidity had no effect on the phenotype of the aid1 mutant. In addition to the partial to complete sterility caused by the defect in pollen development and anther dehiscence, the aid1 mutant showed a delay of 10 to 15 d in heading date and an approximately 40% reduction in tiller numbers; the tiller numbers of the segregated mutant and normal plants in the F4 population were 4.8 and 8.5, respectively.
JA, a multifunctional growth regulator, has been shown to be essential for anther dehiscence in Arabidopsis, and exogenous application of JA has been shown to rescue some of the fertility-related mutations (Sanders et al., 2000
The aid1 Mutant Is Ds Tagged Three different Ds flanking sequences were rescued from mutant plants B4-1-1-1-2, -3, -6, and -8. Database searches indicated that LW602 (accession no. AY429020), LW599 (accession no. AY429019), and LW597 (accession no. AY429018) were rice genomic DNA corresponding to P1-derived artificial chromosome (PAC) P0675A05 (accession no. AP002071), PAC P0015E04 (accession no. AP002069), and bacterial artificial chromosome (BAC) OSJNBa0035I03 (accession no. AP003019), respectively, which all map to the short arm of chromosome 6. To identify which Ds insertion was associated with the spikelet sterility phenotype, a set of primers were synthesized, based on sequences available in the National Center for Biotechnology Information (NCBI) GenBank database, to amplify the region flanking each of the Ds insertion sites (PCR1). PCR2 and PCR3 amplified Ds3' and Ds5' flanking sequence tags (FSTs), respectively (see Table IV for primers used). Assuming that the Ds insertion was in a single-copy gene, using this combination of PCRs, homozygous Ds insertion plants would be PCR1–, PCR2+, and PCR3+; heterozygous Ds insertion plants would be PCR1+, PCR2+, and PCR3+; and Ds null plants would be PCR1+, PCR2–, and PCR3–. These PCR results showed that the Ds insertions mapping to PACs P0675A05 and P0015E04 were present in all 11 F3 plants segregating for spikelet sterility (data not shown), indicating that they were not responsible for the mutant phenotype. The Ds insertion mapping to BAC OSJNBa0035I03 segregated with the mutant phenotype in F3 plants (Table I), indicating that it was responsible for the mutant phenotype.
Cosegregation of this Ds insertion and the mutant phenotype was confirmed in the F4 generation by PCR analyses as described above. All 145 F4 progeny plants of five partially (spikelet) sterile Ds insertion plants (B4-1-1-1-2, -3, -4, -9, and -10) showed partial to complete spikelet sterility and were homozygous for Ds (Ds/Ds), with an average percentage of spikelet sterility of 73% to 92% (Table I). Four plants for the heterozygous Ds insertion segregated at a 1:2:1 (32:57:29;  2 = 0.29, P = 0.87) ratio of homozygous (Ds/Ds), heterozygous (Ds/+), and Ds null (+/+) plants. Observed cosegregation of mutant phenotype and Ds insertion puts the linkage to <0.03 cM, which accounts for less than 11 kb. There were no marked differences in spikelet sterility between Ds/+, +/+ plants, and wild type, while spikelet sterility of Ds/Ds plants ranged from 73% to 78%. The progeny of the Ds null plant B4-1-1-1-12 remained null for Ds and showed a normal phenotype (Table I). This cosegregation relationship was further confirmed by Southern-blot analysis of F4 plants (Fig. 3). Ds null plants were predicted to contain a hybridizing restriction fragment of 3.2 kb. As there is no recognition site for AvrII within Ds (8.5 kb), homozygous Ds insertion plants would have an 11.7-kb hybridizing restriction fragment (Fig. 4B), while heterozygous Ds insertion plants would have both 3.2- and 11.7-kb hybridizing restriction fragments. Hybridization results were as expected (Fig. 3).
AID1 Encodes a Novel Protein with a Single MYB Domain The Ds insertion associated with the mutant phenotype is located at a position corresponding to 133381 nt in BAC OSJNBa0035I03 (Fig. 4, A and B). At the time of the initial analysis, this BAC had not been annotated. Based on gene prediction, expressed sequence tags (ESTs) available in the public domain and reverse transcription (RT)-PCR (data not shown), three putative genes were predicted in a 16-kb region spanning the Ds insertion site (124–140 kb of BAC OSJNBa0035I03). This result is consistent with the annotation now available in The Institute for Genomic Research (TIGR) rice database, but differs from that in NCBI (Fig. 4A). The putative AID1 gene corresponds to TIGR's predicted 3008.t00028 locus and contains five exons (Fig. 4B). As the mutant phenotypes were tightly linked to the Ds insertion and loss of-function of the AID1 gene, and the expression of the gene (3008.t00029 locus) downstream of AID1 was not affected in the aid1 mutant (see below), we concluded that AID1 is the gene involved in the developmental events leading to anther dehiscence in rice. Using RT-PCR and RACE approaches, we cloned the AID1 gene (see "Materials and Methods"; accession no. AY429017). The Ds element in the aid1 mutant was inserted within the 206th codon of the AID1 gene (Fig. 5).
AID1 is predicted to encode a protein of 426 amino acids that contains a single MYB DNA-binding domain at the C terminus. Its MYB domain is closely related to that of telomere-binding proteins, including TRF1 and TRF2 of human and mouse, TRP1 and TBP1 of Arabidopsis, and of single MYB-repeat transcriptional regulators of plants such as MYB1 of P. crispum (Feldbrügge et al., 1997 ), IBP1 of maize (Lugert and Werr, 1994), and BPF1 of P. crispum and Catharanthus roseus (Costa e Silva et al., 1993; van der Fits et al., 2000) as shown in Figure 6. Alignment of the predicted sequence of AID1 and other telomere-binding proteins revealed 36% to 50% sequence identity in the MYB domains. However, there is no similarity outside the MYB domain, suggesting that AID1 is a novel gene.
To determine the expression pattern of the AID1 gene, northern-blot analysis was performed with wild-type RNA using a 5' cDNA probe, but only a very weak signal was detected, indicating that AID1 has low expression. However, AID1 transcripts were detected by RT-PCR in the wild-type panicle as well as leaves, but not in the mutant panicle or leaves. Two transcripts were amplified (Fig. 7). The third exon of the AID1 gene was found to be missing in the smaller transcript, which could result in a premature termination and a nonfunctional protein. RT-PCR also indicated that the expression of the gene downstream of AID1 was not affected in the aid1 mutant (Fig. 7), suggesting that the 3008.t00028 and 3008.t00029 loci are two independent genes.
The aid1 Mutation Affects Events Leading to Anther Dehiscence
We have identified a mutant, aid1, from a Ds-tagged rice-screening population, in which spikelet sterility is the most obvious phenotype. Based on a tight phenotypic segregation with the Ds tag and associated nonexpression of the tagged AID1 gene in the mutant, we conclude that the AID1 gene is important for fertility as well as increased tillering and early flowering. Sterility in the aid1 mutant was mainly caused by a defect in anther dehiscence (type 2 spikelets), although approximately 25% of the spikelet sterility was due to pollen sterility (type 1 spikelets). The failure of anther dehiscence is associated with a defect in the programmed degradation and breakage of anther wall tissues. In the wild type, from the vacuolated microspore stage of anther development, a cavity begins to form because the stomium dissociates from the broken-down septum (Fig. 2, A and B). Further splitting of the stomium, caused by the pressure of swollen pollen grains and the force induced by the inward bending of the locules, results in the release of pollen grains (Fig. 2C). But this process was not observed in the aid1 mutant (Fig. 2, D and E). Degradation of the septum and breakage of the stomium needs both enzymatic lysis of cell wall and mechanical force (Keijzer et al., 1996
The importance of the deposition of lignified cellulose in the secondary wall thickenings of endothecium has been elucidated in Arabidopsis (Steiner-Lange et al., 2003
The fact that a small proportion of the anthers did dehisce normally in the aid1 mutant indicates that AID1 is important, but not absolutely essential, for normal anther development. The exact reason for varying levels of defects in anther development in the mutant is yet to be determined, but it is possible that environmental conditions are important for the induction of these events. For example, a thermosensitive genic male sterile mutant has been shown to be associated with premature programmed cell death of the tapetum (Ku et al., 2003
The involvement of JA in anther dehiscence has been confirmed by isolating genes involved in JA biosynthesis in Arabidopsis (Sanders et al., 2000
MYB domain proteins in plants represent the largest gene family of transcription factors, suggesting their participation in a number of processes. They can be classified into three subfamilies, MYB1R, R2R3-type MYB, and MYB3R, depending on the number of adjacent repeats (1–3) in the MYB domain (for review, see Stracke et al., 2001
The single MYB domain proteins are quite divergent; however, two highly conserved amino acid motifs, SHAQK(Y/F) F (Rose et al., 1999 AID1 is a novel member of the LKDKW type of single MYB domain proteins (Figs. 5 and 6). The aid1 mutation blocked degradation of the septum and splitting of the stomium. It also affected the deposition of starch in pollen grains and the development of fibrous bands in endothecial cells. These results suggest that AID1 regulates programmed cell death as well as deposition of secondary metabolic materials during maturation of pollen grains. Therefore, AID1 is most likely a transcriptional regulator involved in late developmental events concerned with the maturation of pollen grains and anther dehiscence.
The pleiotropic effects observed, such as fewer tillers and delayed flowering, further suggest that AID1 functions as a transcriptional regulator and has a role to play not only in the reproductive stage, but also in the vegetative stage and in the flowering transition. Consistent with this interpretation, AID1 is expressed in both vegetative and reproductive tissues (Fig. 7). Transcription factors can exhibit multiple functions, and a mutation could lead to pleiotropic effects as observed in the rice GAMYB loss-of-function mutation that shows both impaired
The BAC OSJNBa0035I03 harboring the AID1 gene has been anchored to approximately 13.5 cM on chromosome 6. The genetic distance between AID1 and the well-characterized Waxy (Wx) gene located at 8.3 cM on chromosome 6 (Nagato and Yoshimura, 1998
In maize, ms2, which maps to chromosome 9, is also closely linked to Wx (Burnham, 1978 In conclusion, using a transposon Ds-tagged mutant, we have identified a gene in rice, designated AID1, involved in anther development with pleiotropic effects on tillering and flowering time. AID1 is a single MYB DNA-binding domain protein. The conserved motif LKDKW within the MYB domain of AID1 is identical to those of transcription factors TRP1 and TBP1 of Arabidopsis, MYB1 of P. crispum and IBP1 of maize.
Plant Materials and Growth Conditions
All rice (Oryza sativa) lines used in this study were derived from the japonica cv Nipponbare. The male sterility mutant described in this paper was derived from an F2 generation plant (B4-1-1-1) derived from a cross between iAc (pSK300, TT3-26-1) and Ds gene trap (DsG, pSK200, TT6-101-1) transgenic lines described previously by Upadhyaya et al. (2002)
Genomic DNA was extracted from plants using the PureGene nucleic acid isolation kit (Gentra Systems, Minneapolis) according to the manufacturer's instructions. Ds flanking sequences were rescued using the built-in plasmid rescue system (Upadhyaya et al., 2000
PCR primers (Table IV) were designed using the prime program of the Genetics Computer Group (Madison, WI) software suit (Devereux et al., 1984
For Southern-blot analysis, genomic DNA (approximately 10 µg) was extracted from F4 progeny of B4-1-1-1-8 and wild-type plants, digested with AvrII, fractionated on a 0.7% agarose gel, and blotted onto a Hybond-N+ membrane (Amersham Life Science, Buckinghamshire, UK) according to the manufacturer's instructions. A 744-bp genomic region (Fig. 4B) flanking the Ds insertion was amplified with primers OSJN_For and OSJN_Rev (Table IV) and used to prepare a radioactively labeled probe with the Megaprime DNA labeling system (Amersham Life Science) according to the manufacturer's instructions. The membrane was washed at 60°C with 0.1x SSC and 0.1% SDS and visualized by autoradiography using phosphor screens (Molecular Dynamics, Sunnyvale, CA).
Total RNA was extracted from young panicles and leaves by using RNeasy mini kit (Qiagen, Valencia, CA) and treated with DNA-free (Ambion, Austin, TX). Partial 5' cDNA and 3' cDNA (two bands) of the AID1 gene were amplified by primer combination of AID1_fg27_For and AID1_Rev, and OSJN_For and AID1_tigr28_Rev (Fig. 5), respectively, using OneStep RT-PCR kit (Qiagen), according to the manufacturer's instructions. After purification using UltraClean (Mo Bio Laboratories, Solana Beach, CA), bands were cloned into pGEM-T Easy (Promega, Madison, WI) and sequenced using T7 and M13 reverse primers. One 3' cDNA clone was found to contain premature stop codon after sequence analysis. 3' RACE cDNA was isolated with primer AID1_fg27_For2 using SMART RACE cDNA amplification kit (BD Biosciences CLONTECH, Palo Alto, CA), according to the manufacturer's instructions. The rice Suc synthase gene RSs1 (Wang et al., 1992
Investigations on spikelet opening, filament elongation, anther dehiscence, and pollen viability were conducted on the day of anthesis. Spikelets were categorized into three types, based on the behavior of anther dehiscence and pollen viability: (1) no filament elongation—anthers did not elongate out of the spikelet because of no filament elongation and spikelet opening, and showed pollen sterility; (2) anther indehiscence—anthers elongated out of spikelet, but did not dehisce or showed delayed dehiscence, and pollen grains were viable; and (3) anther dehisced—anthers dehisced normally and showed normal pollen viability. Pollen viability was examined by staining with 1% iodine-potassium iodide solution. For wild-type and normal dehisced anthers of the aid1 mutant (type 3), pollen grains were directly collected on a slide by shaking dehiscing anthers. For indehiscence anthers of the aid1 mutant (types 1 and 2), pollen grains were manually squeezed onto a slide. Observations were immediately made by microscopic investigation following the addition of a drop of 1% iodine-potassium iodide solution. Ten spikelets were checked for each type of spikelet.
For scanning electron micrograph observation, anthers were collected from a spikelet at its peak stage of anthesis and processed essentially as described by Keijzer et al. (1996)
Plants of wild type and the aid1 mutant were tagged immediately prior to booting. Tissue paper was wrapped outside the leaf sheath of the flag leaf. Three panicles of each plant were given one of the following treatments: (1) no treatment (control); (2) water; and (3) 500 µM MeJA (Sigma-Aldrich, St. Louis). The three treatments were applied twice daily (morning and dusk) by applying 2 mL of water or MeJA solution to the tissue paper using a syringe. The treatment lasted for 15 d. Each treatment included five plants (replications). Fertile and sterile spikelets were counted separately for each tagged panicle to determine the percentage of spikelet sterility. Sequence data from this article have been deposited with the EMBL/GenBank data libraries under accession numbers AY429017, AY429020, AY429019, and AY429018.
The authors thank Celia Miller, Shu-Ting Pan, Shamsul Hoque, and Kathryn Smith for their invaluable technical assistance. We also thank Drs. John Watson, Tony Pryor, Ming-Bo Wang, and Andrew Eamens for their critical reading and discussion. We are grateful to our collaborators, Chellian Santhoshkumar and Kottaram K. Narayanan, for providing pSK200 and pSK300 constructs. Received February 25, 2004; returned for revision April 18, 2004; accepted April 21, 2004.
1 This work was supported in part by GrainGene, Australia.  Article, publication date, and citation information can be found at www.plantphysiol.org/cgi/doi/10.1104/pp.104.041459. * Corresponding author; e-mail narayana.upadhyaya{at}csiro.au; fax 61–2–6246–5000.
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-amylase in aleurone, but also in pollen development (Kaneko et al., 2004
