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• Photoperiod-dependent male fertility is a critical enabler of modern hybrid breeding. A MYB transcription factor, CSA, is a key regulator of sugar partitioning in rice anthers, disruption of which causes photoperiod-sensitive male sterility. However, little is known about the molecular mechanisms governing plant fertility in response to photoperiod. • Here, we have obtained another rice photoperiod-sensitive male sterile mutant, csa2, which exhibits semi-sterility under long-day (LD) conditions, with normal fertility under short-day (SD) conditions. CSA2 specifically expressed in anthers, and here is shown to be indispensable for sugar partitioning to anthers under LD conditions. • The CSA2 protein can restore the fertility of csa mutants under SD conditions when expressed in a CSA-specific pattern, indicating that the two proteins share common downstream regulatory targets. Transcriptomic analyses also reveal discrete regulatory targets in anthers. Furthermore, the regulatory role of CSA2 in sugar transport was influenced by the photoperiod conditions during floral initiation, not simply during anther development. • Collectively, we propose that rice evolved at least two MYB proteins, CSA2 and CSA, that regulate sugar transport in anthers under LD and SD conditions, respectively. This finding provides insight into the molecular mechanisms that regulate male fertility in response to photoperiod.

Background Decreased spikelet fertility is often responsible for reduction in grain yield in rice ( Oryza sativa L.). In this study, two varieties with different levels of heat tolerance, Liangyoupeijiu (LYPJ, heat susceptible) and Shanyou63 (SY63, heat tolerant) were subjected to two temperature treatments for 28 days during the panicle initiation stage in temperature/relative humidity-controlled greenhouses: high temperature (HT; 37/27 °C; day/night) and control temperature (CK; 31/27 °C; day/night) to investigate changes in anther development under HT during panicle initiation and their relationship with spikelet fertility. Results HT significantly decreased the grain yield of LYPJ by decreasing the number of spikelets per panicle and seed setting percentage. In addition, HT produced minor adverse effects in SY63. The decreased spikelet fertility was primarily attributed to decreased pollen viability and anther dehiscence, as well as poor pollen shedding of the anthers of LYPJ under HT. HT resulted in abnormal anther development (fewer vacuolated microspores, un-degraded tapetum, unevenly distributed Ubisch bodies) and malformation of pollen (obscure outline of the pollen exine with a collapsed bacula, disordered tectum, and no nexine of the pollen walls, uneven sporopollenin deposition on the surface of pollen grains) in LYPJ, which may have lowered pollen viability. Additionally, HT produced a compact knitted anther cuticle structure of the epidermis, an un-degraded septum, a thickened anther wall, unevenly distributed Ubisch bodies, and inhibition of the confluent locule, and these malformed structures may be partially responsible for the decreased anther dehiscence rate and reduced pollen shedding of the anthers in LYPJ. In contrast, the anther wall and pollen development of SY63 were not substantially changed under HT. Conclusions Our results suggest that disturbed anther walls and pollen development are responsible for the reduced spikelet fertility and grain yield of the tested heat susceptible variety, and noninvasive anthers and pollen formation in response to HT were associated with improved heat tolerance.

• Global warming has reduced the productivity of many field-grown crops, as the effects of high temperatures can lead to male sterility in such plants. Genetic regulation of the high temperature (HT) response in the major crop cotton is poorly understood. • We determined the functionality and transcriptomes of the anthers of 218 cotton accessions grown under HT stress. By analyzing transcriptome divergence and implementing a genome-wide association study (GWAS), we identified three thermal tolerance associated loci which contained 75 protein coding genes and 27 long noncoding RNAs, and provided expression quantitative trait loci (eQTLs) for 13 132 transcripts. • A transcriptome-wide association study (TWAS) confirmed six causal elements for the HT response (three genes overlapped with the GWAS results) which are involved in protein kinase activity. The most susceptible gene, GhHRK1, was confirmed to be a previously uncharacterized negative regulator of the HT response in both cotton and Arabidopsis. • These functional variants provide a new understanding of the genetic basis for HT tolerance in male reproductive organs.

• Degeneration of apical spikelets and reduced panicle fertility are common reasons for low seed-setting rate in rice (Oryza sativa). However, little is known about the underlying molecular mechanisms. • Here, we report a novel degenerated panicle and partial sterility 1 (dps1) mutant that showed panicle apical degeneration and reduced fertility in middle spikelets. dps1 plants were characterized by small whitish anthers with altered cuticle morphology and absence of pollen grains. Amounts of cuticular wax and cutin were significantly reduced in dps1 anthers. Panicles of dps1 plants showed an accumulation of reactive oxygen species (ROS), lower antioxidant activity, and increased programmed cell death. • Map-based cloning revealed that DPS1 encodes a mitochondrial-localized protein containing a cystathionine β-synthase domain that showed the highest expression in panicles and anthers. DPS1 physically interacted with mitochondrial thioredoxin proteins Trx1 and Trx20, and it participated in ROS scavenging. Global gene expression analysis in dps1 revealed that biological processes related to fatty acid metabolism and ROS homeostasis were significantly affected, and the expression of key genes involved in wax and cutin biosynthesis were downregulated. • These results suggest that DPS1 plays a vital role in regulating ROS homeostasis, anther cuticle formation, and panicle development in rice.

Litchi ( Litchi chinensis Sonn.), an important fruit originating in China, has received increasing attention for improving breeding efficiency in recent years due to the growing adoption of hybridization. It is a monoecious plant and predominantly relies on cross-pollination for seed production. Thus, enhancing pollen quality is essential for fruit set, which is closely associated with the developmental stages of male flowers. In this study, anthers of the litchi cv. ‘D13’ at five late developmental stages (half-exposed stage, fully-exposed stage, filament-elongated stage, anthesis, and browning stage) were investigated using paraffin sectioning, staining, in vitro germination, and controlled pollination techniques. Cytological characteristics of pollen were evaluated using paraffin sectioning, while pollen viability was assessed by I 2 -KI staining, and the germination rate was determined through in vitro culture on sucrose media. Furthermore, pollen collected from these five stages was used for controlled pollination of the male-sterile line ‘MS1’. The results indicated that pollen collected at the filament-elongated stage and anthesis exhibited higher viability and germination potential, which resulted in improved fruit set in the male-sterile line ‘MS1’. The study identified the filament-elongated stage and anthesis as optimal stages for pollen collection in litchi.

Key message Anther development and dehiscence is considered from an evolutionary perspective to identify drivers for differentiation, functional conservation and to identify key questions for future male reproduction research. Development of viable pollen and its timely release from the anther are essential for fertilisation of angiosperm flowers. The formation and subsequent dehiscence of the anther are under tight regulatory control, and these processes are remarkably conserved throughout the diverse families of the angiosperm clade. Anther development is a complex process, which requires timely formation and communication between the multiple somatic anther cell layers (the epidermis, endothecium, middle layer and tapetum) and the developing pollen. These layers go through regulated development and selective degeneration to facilitate the formation and ultimate release of the pollen grains. Insight into the evolution and divergence of anther development and dehiscence, especially between monocots and dicots, is driving greater understanding of the male reproductive process and increased, resilient crop yields. This review focuses on anther structure from an evolutionary perspective by highlighting their diversity across plant species. We summarise new findings that illustrate the complexities of anther development and evaluate how they challenge established models of anther form and function, and how they may help to deliver future sustainable crop yields.

Main ConclusionThe characteristics of sorghum anthers at 18 classified developmental stages provide an important reference for future studies on sorghum reproductive biology and abiotic stress tolerance of sorghum pollen.Sorghum (Sorghum bicolor L. Moench) is the fifth-most important cereal crop in the world. It has relatively high resilience to drought and high temperature stresses during vegetative growing stages comparing to other major cereal crops. However, like other cereal crops, the sensitivity of male organ to heat and drought can severely depress sorghum yield due to reduced fertility and pollination efficiency if the stress occurs at the reproductive stage. Identification of the most vulnerable stages and the genes and genetic networks that differentially regulate the abiotic stress responses during anther development are two critical prerequisites for targeted molecular trait selection and for enhanced environmentally resilient sorghum in breeding using a variety of genetic modification strategies. However, in sorghum, anther developmental stages have not been determined. The distinctive cellular characteristics associated with anther development have not been well examined. Lack of such critical information is a major obstacle in the studies of anther and pollen development in sorghum. In this study, we examined the morphological changes of sorghum anthers at cellular level during entire male organ development processes using a modified high-throughput imaging variable pressure scanning electron microscopy and traditional light microscopy methods. We divided sorghum anther development into 18 distinctive stages and provided detailed description of the morphological changes in sorghum anthers for each stage. The findings of this study will serve as an important reference for future studies focusing on sorghum physiology, reproductive biology, genetics, and genomics.

Significance Germ cells (GCs) (i.e., the cells that are committed to meiosis and gametogenesis) are key carriers for eukaryotes to complete their life cycle, transmitting their genetic information from one generation to the next while generating variations to integrate environmental changes. Compared to what has been known in animals, very little is known about how the GCs in plants are segregated from somatic cells. This work demonstrates that auxin is a key factor guiding GC specification in Arabidopsis anthers. Local auxin biosynthesis interacts with the transcription of SPOROCYTELESS/NOZZLE and a progressive GC specification itself to form a dynamic feedback circuit that ensures the completion of GC specification.

Cold stress is a serious threat to subtropical crop pollen development and induces yield decline. N6-methyladenosine (m 6 A) is the most frequent mRNA modification and plays multiple physiological functions in plant development. However, whether m 6 A regulates pollen development is unclear, and its putative role in cold stress response remains unknown. Here, we observed that moderate low-temperature (MLT) stress induced pollen abortion in tomato. This phenotype was caused by disruption of tapetum development and pollen exine formation, accompanied by reduced m 6 A levels in tomato anther. Analysis of m 6 A-seq data revealed 1,805 transcripts displayed reduced m 6 A levels and 978 transcripts showed elevated m 6 A levels in MLT-stressed anthers compared with those in anthers under normal temperature. These differentially m 6 A enriched transcripts under MLT stress were mainly related to lipid metabolism, adenosine triphosphatase (ATPase) activity, and ATP-binding pathways. An ATP-binding transcript, SlABCG31 , had significantly upregulated m 6 A modification levels, which was inversely correlated to the dramatically downregulated expression level. These changes correlated with higher abscisic acid (ABA) levels in anthers and disrupted pollen wall formation under low-temperature stress. Our findings characterized m 6 A as a novel layer of complexity in gene expression regulation and established a molecular link between m 6 A methylation and tomato anther development under low-temperature conditions.