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Frontiers in Plant Science

Plant Genetics and Genomics

Original Research ARTICLE

Front. Plant Sci., 05 February 2018 | https://doi.org/10.3389/fpls.2018.00091

Cold-Induced Male Meiotic Restitution in Arabidopsis thaliana Is Not Mediated by GA-DELLA Signaling

  • 1Department of Plant Production, Faculty of Bioscience Engineering, University of Ghent, Ghent, Belgium
  • 2School of Integrative Plant Science, Cornell University, Ithaca, NY, United States

Short periods of cold stress induce male meiotic restitution and diploid pollen formation in Arabidopsis thaliana by specifically interfering with male meiotic cytokinesis. Similar alterations in male meiotic cell division and gametophytic ploidy stability occur when gibberellic acid (GA) signaling is perturbed in developing anthers. In this study, we found that exogenous application of GA primarily induces second division restitution (SDR)-type pollen in Arabidopsis, similar to what cold does. Driven by the close similarity in cellular defects, we tested the hypothesis that cold-induced meiotic restitution is mediated by GA-DELLA signaling. Using a combination of chemical, genetic and cytological approaches, however, we found that both exogenously and endogenously altered GA signaling do not affect the cold sensitivity of male meiotic cytokinesis. Moreover, in vivo localization study using a GFP-tagged version of RGA protein revealed that cold does not affect the expression pattern and abundance of DELLA in Arabidopsis anthers at tetrad stage. Expression study found that transcript of RGA appears enhanced in cold-stressed young flower buds. Since our previous work demonstrated that loss of function of DELLA causes irregular male meiotic cytokinesis, we here conclude that cold-induced meiotic restitution is not mediated by DELLA-dependent GA signaling.

Introduction

The production of viable haploid gametes is vital for the fertility and ploidy stability of flowering plants. Under certain conditions, plants may produce unreduced male gametes through incomplete meiotic cell division, a phenomenon termed ‘meiotic restitution’ or ‘meiotic non-reduction’ (Adams and Wendel, 2005;Mason and Pires, 2015). Meiotic restitution can be induced by omission of meiotic cell cycles, defective spindle organization, and/or irregular meiotic cytokinesis (De Storme and Geelen, 2013a; De Storme and Mason, 2014). Meiotic restitution can be classified into either FDR or SDR according to the genetic make-up of the yielded unreduced gametes (Kohler et al., 2010). In FDR-type meiotic restitution, homologous chromosomes fail to separate whereas sister chromatids successfully disjoin from each other. As a result, FDR-type unreduced gametes maintain parental heterozygosity in genomic regions where meiotic recombination occurs rarely. In contrast, in SDR-type meiotic restitution, homologous chromosomes segregate but sister chromatids are grouped into unreduced gametes, which typically leads to heterozygosity at recombination-free chromatin regions (De Storme and Geelen, 2013b). Meiotic restitution and associated formation of unreduced gametes can be induced either through specific genetic defects or by temperature stress. In Arabidopsis thaliana, for example, cold stress causes male meiotic restitution (primarily SDR-type) by specifically disrupting the organization of RMAs at the end of male meiosis, consequently resulting in incomplete meiotic cytokinesis and unreduced gamete formation (De Storme et al., 2012).

The phytohormone GA regulates multiple development processes during male reproduction in plants (Plackett et al., 2011; Kwon and Paek, 2016; Plackett and Wilson, 2016). In most cases, GA regulates plant development by suppressing the activity of DELLAs, a family of transcriptional repressors that operate as negative regulators of GA signaling (Sun, 2010). DELLA proteins negatively regulate the expression of GA-signaling downstream genes and consequently inhibit plant growth and development (Xu et al., 2014). At the molecular level, GA suppresses DELLA activity by promoting DELLA protein destabilization through 26S proteasome. GA-dependent DELLA degradation relies on activity of two F-box proteins SLEEPY1 (SLY1) and SNEEZY (SNE), and the phosphorylation status of DELLA itself (Ariizumi et al., 2011; Ariizumi and Steber, 2011; Qin et al., 2014). In Arabidopsis, there are five DELLA homologs; i.e., RGA, GAI, RGL1, RGL2, and RGL3.

During male reproductive development, programmed degradation of the tapetal cell layer during male gametogenesis and pollen maturation strongly relies on balanced GA signaling and alterations herein lead to defects in tapetal disintegration, eventually causing abortion of microspores or mature pollen grains (Cheng et al., 2004; Aya et al., 2009; Plackett et al., 2014). GA signaling also plays a role in one or more process of microsporogenesis; e.g., meiotic cytokinesis. Ectopic activation of GA signaling in Arabidopsis meiosis-stage anthers, either via exogenous GA treatment or in the DELLA rga-/- gai-/- mutant background, was recently found to cause defects in male meiotic cell wall formation, leading to ectopic events of male meiotic restitution and associated formation of unreduced (2n) spores (Liu et al., 2017b). These findings suggest that balanced GA levels and associated GA signaling in the tapetal cell layer contributes to the progression of male meiotic cell division and is required for gametophytic ploidy stability (i.e., haploid spores). As the cellular mechanism of GA-induced male meiotic restitution highly mimics the meiotic alterations induced by cold (i.e., defects in RMA formation at telophase II and binuclear spore formation), we hypothesized that cold-induced male meiotic restitution in Arabidopsis could be mediated by GA-DELLA signaling. For this hypothesis to be correct, bioactive GA levels would need to increase in response to cold in the Arabidopsis anthers, in contrast to a general notion that low temperature stress causes a reduction in GA level and thereby promoting DELLA accumulation (Colebrook et al., 2014). In Arabidopsis seedlings, cold-induced accumulation of DELLAs is achieved by enhancing the catabolism of endogenous bioactive GA, through the rapid transcriptional activation of DREB1b/C-repeat/DRE Binding Factor1 (CBF1), a positive regulator of GA-deactivating GA2OX2 (Achard et al., 2008). Similarly, in rice, low temperature reduces the level of endogenous bioactive GA in developing anthers, where the expression of the GA biosynthesis gene OsGA3OX1 is down-regulated upon cold stress, whereas the GA signaling repressor SLR1/DELLA and its upstream cold-responsive factor CBF1 are up-regulated (Sakata et al., 2014).

Here, in this study, we test the hypothesis that cold-induced meiotic restitution in Arabidopsis male sporogenesis is mediated by alterations in anther GA signaling. In support of our hypothesis, we found that exogenous GA treatment of Arabidopsis primarily induces SDR-type unreduced gametes, in a similar rate and manner as when plants are exposed to cold. However, contrary to the assumption that GA may mediate the cold response of male meiosis, our data indicated that the cold sensitivity of male sporogenesis does not rely on the DELLA-dependent GA signaling. In addition, we found that cold stress does not reduce RGA abundance in the young developing anthers. Together these findings indicate that cold-induced male meiotic restitution in Arabidopsis is not mediated by GA-DELLA signaling.

Materials and Methods

Plant Materials and Growth Conditions

Arabidopsis (Arabidopsis thaliana) Columbia-0 and Landsberg erecta (Ler) accessions were obtained from the Nottingham Arabidopsis Stock Centre. The GA-insensitive mutant gai was kindly shared by Patrick Achard. The DELLA double mutant rga-24 gai-t6 was previously described (Achard et al., 2006) and were kindly provided by Patrick Achard and Nicholas Harberd. The fluorescent tagged lines (FTLs) in the qrt1-2-/- background (FTL1313 and FTL3332), used for genotyping unreduced gametes, were described earlier (Francis et al., 2007; Berchowitz and Copenhaver, 2008). FTL1313 marker is physically located at 11.2 cM on chromosome 1, and FTL3332 is positioned at 10.43 cM on chromosome 31 (Supplementary Figure S1A) (Francis et al., 2007; Singh et al., 2017). The pRGA::GFP-4k5a>; ; Sin13b). Meiotic res--alintitself (Ariizumi et h3b). Meiotic r0aver, 2008). FTL1313 m response to c-4h3b). Me4otic r0aver,42008). F(FTL131k5response to 4-4h3b). p Me4otic rk5r,42008)-name="h3m ). pu.4kgtic  the Notti" data-html="True">

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(1997).4vk5 pt50P4uB4k5kw4 GAIk5g(g-t/Tdefine4kwe5e5e55e5 pathwayvtkatw4eg4k4v0lykpe14a5p4 gi5O,45hk5kkpesponse4v;pt5G-t/s;Dkv.k5g(41, 3194p0p13;3205u54sik5fbc44p1/gadi11.23.31945isiskg4Tfltnflnfltnfltisisisisisisisisvr444k5p444k5p44444k5p444k5p444k5pH44k5p444k5p44(r4k544k5p444k5p4u4k544k5p444k5p4u4k544k5p4449389651yft5t-kmtnSn;););)>v4k1bgf1yettge-su4k0kgt52e0e055047474t4t4t4v4747u44p1/gadi11.23.3194yft5t-kmtnSn;););)lrom 5kw N,dPf2-k547t-kmtnSvk54kt kr5kvr444(hft5t-kmtnSn;);););mm5oky55k55/ct0554k5kmtnSvk550P4uB4k5kw4+GAI+g-t/+define45a+e5e5e55e5+pathway/tkat+4eg4k4v0lympe14a5p4+gi5O,45hk5kmpesponse44n.vm4l5k4skr4r4r=G-t/s+Dkv4n.vm4l5k4kkm>v4kPeng+Jvsisisisikkm>v4kCapol+Pvsisisisikkm>v4kRichards5p+Evsisisisikkm>v4kK5e5+Kp+Evsisisisikkm>v4kCowlin5+Rv+Jvsisisisikkm>v4kkurph+G.+Pvsisisisi444k5c4k4kw4k5p441997sisisisi4m>v5p441p0isisi555Ht3194-3205yft5t-kmtnSn;););)n4ky3ky3i 5k5g(5t5isiskg.4k5t4km44/hkk]440laht5kg45t50v;pt54en41)I5t/3i 5k5kvu4kvT4T4T4TfltnflnfltnfltB4k5kw4k5p4434yft5unn344k5(5tml)ckptthkA/-RihkFerguso5hkA/-CihkPowers--S.-J/hkWithoo-KohlihkA/hkPhillips--A/-L/hkWilso5hkZ. 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A/ (p16504sisp1C;Gt5O,45hk5k4knkk);)nt repro5u4k4kw,sisp1D; Htvpt5Annual ml)nt Rev44k4k5g, Vok. 49hk di2S.-Mep44)nt (Hobok4khkNJ: JohnkWiley4sisisisi Sons), 323p0p13;358.k5siskg4Tfltnflnfltnfltisisisisisisisisvr4;);mm5ot-kmtnSn;);););mm5oky55k55/ct0554k5kmtnSGt5O,45hk5k45ank+);)nt+repro5u4k4kw/ct0554kskr4r4r=Annual+ml)k5+Rev44k4/ct0554kkkm>v4kml)ckpttk4k++Rv++G.vm4l5k4kkm>v4kank+Wilso5+Z.kk4kvm4l5k4444k5c4k4kw4k5p445t16sisisisi4m>v5p449sisisisi555Ht323-358yft5t-kmtnSn;););)n4ky3ky3i 5k5g(5t5isiskg.4k5t4km44/hkk]440laht5kg45t50v;pt54en41)I5t/3i 5k5kvu4kvT4T4T4TfltnflnfltnfltB4k5kw4k5p4437yft5unn374k5(5tQikhkQ/hkWit5--W/hkGuo2X/hkYue--J/--Huit5--Y/--Xu--X/-- 5kak. 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B/hkank Jot/s--A/-Mi (p17).4In vivo gi5O,45hk5kkgradi4k5s visue55z1bg5kkrapidlyk0long4k4e5 tissue4v;pt5Nat. 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(p14504Re5u4k4kw of gi5O,45hk5kgbylow temppritur/Tdisrup5s poll 5vdev0lopm4k5g5kkricev;pt5ml)k5kPh54k.k5g(164, 2t11sis2t13;2019044t4k5fbc44p4/ppc443.2344015isiskg4Tfltnflnfltnfltisisisisisisisisvr444k5p444k5p44444k5p444k5p444k5pH44k5p444k5p44(r4k544k5p444k5p4u4k544k5p444k5p4u4k544k5p44424569847yft5t-kmtnSn;););)>v4k1bgf1yettge-su4k0kgt52e0e055047474t4t4t4v4747u44p4/ppc443.234401yft5t-kmtnSn;););)lrom 5kw N,dPf2-k547t-kmtnSvk54kt kr5kvr444(hft5t-kmtnSn;);););mm5oky55k55/ct0554k5kmtnSRe5u4k4kw+u4/gi5O,45hk5k+by5low+temppritur/+disrup5s/poll 5ndev0lopm4k55g5rice4n./ct0554kskr4r4r=ml)k5+ne54k4n./ct0554kkkm>v4kSakata+Tvsisisisikkm>v4kOdaim>sisisisikkm>v4kTsunaga+Yvsisisisikkm>v4k444mura+Hvsisisisikkm>v4kKawagishi-KobayashikMvsisisisikkm>v4kAya+Kpsisisisi444k5c4k4kw4k5p445t14p0isisi4m>v5p4164p0isisi555Ht5t11-5t19yft5t-kmtnSn;););)n4ky3ky3i 5k5g(5t5isiskg.4k5t4km44/hkk]440laht5kg45t50v;pt54en41)I5t/3i 5k5kvu4kvT4T4T4TfltnflnfltnfltB4k5kw4k5p4440yft5unn40yt5i5tSalomp0isE9;hkP. A/hkBom4k5/s--K/--Fitz--J/--Laik4e4khkR.-A/-E/--WarthmankhkN/--Y)k5--L/-- 5kak. 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