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Research Progress on Maize Dwarf Genes
LIJintao, SHEKuijun, WURui, YANGGuohu
Chin Agric Sci Bull ›› 2025, Vol. 41 ›› Issue (32) : 9-16.
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Abbreviation (ISO4): Chin Agric Sci Bull
Editor in chief: Yulong YIN
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Research Progress on Maize Dwarf Genes
Maize is the crop with the largest planting area in China, playing a crucial role in safeguarding national food security. Dwarfing breeding is a core approach to break the bottleneck of maize yield per unit area by optimizing plant architecture and increasing planting density. This paper systematically reviews the research progress on maize dwarf genes, with a focus on clarifying the biological significance of maize plant height traits, the practices of dwarf genetic breeding, and the regulatory mechanisms of plant hormones on plant height, while proposing future research directions. The results show that: (1) maize plant height is co-regulated by the number of internodes and internode length. Dwarf plants can reduce lodging risk by shortening internode length, optimize canopy structure, and improve light energy use efficiency and adaptability to dense planting, but it is necessary to coordinate the relationship between dwarfing and yield traits. (2) Maize dwarf genetics is divided into two major systems: single-gene and multi-gene. In the single-gene system, the br2 gene has the clearest molecular mechanism—it inhibits the elongation of stem cells, reducing stem length by 40% to 50% compared with the wild type, with a more significant effect on internodes below the ear position—and it is the most widely used major gene at present. The multi-gene system can avoid the defect of pleiotropy by accumulating minor-effect genes, and varieties such as 'Aidan 268' that balance dwarfing and high yield have been bred. (3) Gibberellin (GA), brassinosteroid (BR), and auxin (IAA) are the core hormones regulating plant height: mutations in GA synthesis-related genes (d1, an1) or signal genes (d8, d9) lead to dwarfing, loss of function of BR synthesis genes (brd1, na2) or signal genes (ZmBRI1a) causes stunted plants, and abnormal function of the IAA polar transport gene (br2) results in dwarfing of lower stem nodes. Currently, maize dwarf breeding has problems such as a relatively small number of applicable genes (more than 60 dwarf genes have been discovered, and about 40 have been cloned), genetic linkage drag restricting the coordination of traits, and insufficient functional verification of novel dwarf genes (such as the mapped genes K718d and d8227). In the future, it is necessary to explore medium dwarf genes suitable for dense planting, use genome-wide selection technology to aggregate multiple genes, and integrate phenomics with artificial intelligence to screen for ideal plant architecture, so as to breed maize varieties with the characters of dwarf stalks for lodging resistance, dense planting for high yield, wide adaptability and easy mechanical harvesting, and provide support for the sustainable development of the maize industry.
maize / plant height / dwarf gene / genetic breeding / plant hormone / plant height regulation / br2 gene
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Brassinosteroids (BRs) are plant hormones involved in many developmental processes as well as in plant-environment interactions. Their role was investigated in this study through the analysis of lilliputian1-1 (lil1-1), a dwarf mutant impaired in BR biosynthesis in maize (Zea mays).We isolated lil1-1 through transposon tagging in maize. The action of lil1 was investigated through morphological and genetic analysis. Moreover, by comparing lil1-1 mutant and wild-type individuals grown under drought stress, the effect of BR reduction on the response to drought stress was examined.lil1-1 is a novel allele of the brassinosteroid-deficient dwarf1 (brd1) gene, encoding a brassinosteroid C-6 oxidase. We show in this study that lil1 is epistatic to nana plant1 (na1), a BR gene involved in earlier steps of the pathway. The lill-1 mutation causes alteration in the root gravitropic response, leaf epidermal cell density, epicuticular wax deposition and seedling adaptation to water scarcity conditions.Lack of active BR molecules in maize causes a pleiotropic effect on plant development and improves seedling tolerance of drought. BR-deficient maize mutants can thus be instrumental in unravelling novel mechanisms on which plant adaptations to abiotic stress are based.
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Structural variation in plant genomes is a significant driver of phenotypic variability in traits important for the domestication and productivity of crop species. Among these are traits that depend on functional meristems, populations of stem cells maintained by the CLAVATA-WUSCHEL (CLV-WUS) negative feedback-loop that controls the expression of the WUS homeobox transcription factor. WUS function and impact on maize development and yield remain largely unexplored. Here we show that the maize dominant Barren inflorescence3 (Bif3) mutant harbors a tandem duplicated copy of the ZmWUS1 gene, ZmWUS1-B, whose novel promoter enhances transcription in a ring-like pattern. Overexpression of ZmWUS1-B is due to multimerized binding sites for type-B RESPONSE REGULATORs (RRs), key transcription factors in cytokinin signaling. Hypersensitivity to cytokinin causes stem cell overproliferation and major rearrangements of Bif3 inflorescence meristems, leading to the formation of ball-shaped ears and severely affecting productivity. These findings establish ZmWUS1 as an essential meristem size regulator in maize and highlight the striking effect of cis-regulatory variation on a key developmental program.
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Brassinosteroids (BRs) are plant hormones that regulate growth and development. They share structural similarities with animal steroids, which are decisive factors of sex determination. BRs are known to regulate morphogenesis and environmental stress responses, but their involvement in sex determination in plants has been only speculative. We show that BRs control sex determination in maize revealed through characterization of the classical dwarf mutant nana plant1 (na1), which also feminizes male flowers. na1 plants carry a loss-of-function mutation in a DET2 homolog--a gene in the BR biosynthetic pathway. The mutant accumulates the DET2-specific substrate (24R)-24-methylcholest-4-en-3-one with a concomitant decrease of downstream BR metabolites. Treatment of wild-type maize plants with BR biosynthesis inhibitors completely mimicked both dwarf and tasselseed phenotypes of na1 mutants. Tissue-specific na1 expression in anthers throughout their development supports the hypothesis that BRs promote masculinity of the male inflorescence. These findings suggest that, in the monoecious plant maize, BRs have been coopted to perform a sex determination function not found in plants with bisexual flowers.
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Development in higher plants depends on the activity of meristems,formative regions that continuously initiate new organs at their flanks. Meristems must maintain a balance between stem cell renewal and organ initiation. In fasciated mutants, organ initiation fails to keep pace with meristem proliferation. The thick tassel dwarf1 (td1)mutation of maize affects both male and female inflorescence development. The female inflorescence, which results in the ear, is fasciated, with extra rows of kernels. The male inflorescence, or tassel, shows an increase in spikelet density. Floral meristems are also affected in td1 mutants; for example, male florets have an increase in stamen number. These results suggest that td1 functions in the inflorescence to limit meristem size. In addition, td1 mutants are slightly shorter than normal siblings,indicating that td1 also plays a role in vegetative development. td1 encodes a leucine-rich repeat receptor-like kinase (LRR-RLK) that is a putative ortholog of the Arabidopsis CLAVATA1 protein. These results complement previous work showing that fasciated ear2 encodes a CLAVATA2-like protein, and suggest that the CLAVATA signaling pathway is conserved in monocots. td1 maps in the vicinity of quantitative trait loci that affect seed row number, spikelet density and plant height. We discuss the possible selection pressures on td1 during maize domestication.
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In maize (Zea mays L.), as in other grass species, stem elongation occurs during growth and most noticeably upon the transition to flowering. Genes that reduce stem elongation have been important to reduce stem breakage, or lodging. Stem elongation has been mediated by dwarf and brachytic/brevis plant mutants that affect giberellic acid and auxin pathways, respectively. Maize brevis plant1 (bv1) mutants, first identified over 80 years ago, strongly resemble brachytic2 mutants that have shortened internodes, short internode cells, and are deficient in auxin transport. Here, we characterized two novel bv1 maize mutants. We found that an inositol polyphosphate 5-phosphatase orthologue of the rice gene dwarf50 was the molecular basis for the bv1 phenotype, implicating auxin-mediated inositol polyphosphate and/or phosphoinositide signalling in stem elongation. We suggest that auxin-mediated internode elongation involves processes that also contribute to stem gravitropism. Genes misregulated in bv1 mutants included genes important for cell wall synthesis, transmembrane transport, and cytoskeletal function. Mutant and wild-type plants were indistinguishable early in development, responded similarly to changes in light quality, had unaltered flowering times, and had normal flower development. These attributes suggest that breeding could utilize bv1 alleles to increase crop grain yields.© The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology.
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Auxin plays a fundamental role in organogenesis in plants. Multiple pathways for auxin biosynthesis have been proposed, but none of the predicted pathways are completely understood. Here, we report the positional cloning and characterization of the vanishing tassel2 (vt2) gene of maize (Zea mays). Phylogenetic analyses indicate that vt2 is a co-ortholog of TRYPTOPHAN AMINOTRANSFERASE OF ARABIDOPSIS1 (TAA1), which converts Trp to indole-3-pyruvic acid in one of four hypothesized Trp-dependent auxin biosynthesis pathways. Unlike single mutations in TAA1, which cause subtle morphological phenotypes in Arabidopsis thaliana, vt2 mutants have dramatic effects on vegetative and reproductive development. vt2 mutants share many similarities with sparse inflorescence1 (spi1) mutants in maize. spi1 is proposed to encode an enzyme in the tryptamine pathway for Trp-dependent auxin biosynthesis, although this biochemical activity has recently been questioned. Surprisingly, spi1 vt2 double mutants had only a slightly more severe phenotype than vt2 single mutants. Furthermore, both spi1 and vt2 single mutants exhibited a reduction in free auxin levels, but the spi1 vt2 double mutants did not have a further reduction compared with vt2 single mutants. Therefore, both spi1 and vt2 function in auxin biosynthesis in maize, possibly in the same pathway rather than independently as previously proposed.
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The maize crinkly4 (cr4) mutation affects leaf epidermis differentiation such that cell size and morphology are altered, and surface functions are compromised, allowing graft-like fusions between organs. In the seed, loss of cr4 inhibits aleurone formation in a pattern that reflects the normal progression of differentiation over the developing endosperm surface. The cr4 gene was isolated by transposon tagging and found to encode a putative receptor kinase. The extracellular domain contains a cysteine-rich region similar to the ligand binding domain in mammalian tumor necrosis factor receptors (TNFRs) and seven copies of a previously unknown 39-amino acid repeat. The results suggest a role for cr4 in a differentiation signal.
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Bioactive gibberellins (GAs) are tetracyclic diterpenoid plant hormones that promote important processes of plant growth and development, such as seed germination, growth through elongation, and floral transition. Thus, mutant plants that are affected in GA biosynthesis or signalling exhibit altered seed germination and, at the adult stage, are dwarf and dark green and also show delayed flowering. The components of the GA metabolism and signalling pathways are reviewed here and recent findings regarding the regulation and possible mode of action of DELLA proteins are discussed.
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Bacteriophages have been proposed as an alternative to pesticides to kill bacterial pathogens of crops. However, the efficacy of phage biocontrol is variable and poorly understood in natural rhizosphere microbiomes. We studied biocontrol efficacy of different phage combinations on Ralstonia solanacearum infection in tomato. Increasing the number of phages in combinations decreased the incidence of disease by up to 80% in greenhouse and field experiments during a single crop season. The decreased incidence of disease was explained by a reduction in pathogen density and the selection for phage-resistant but slow-growing pathogen strains, together with enrichment for bacterial species that were antagonistic toward R. solanacearum. Phage treatment did not affect the existing rhizosphere microbiota. Specific phage combinations have potential as precision tools to control plant pathogenic bacteria.
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The data presented here show that, in cultures of type 1 astrocytes obtained from the hypothalamus of neonatal female rat, 17β‐oestradiol is able to increase both the mRNA and the protein levels of basic fibroblast growth factor (bFGF). In particular, after 24 h of exposure to 17β‐oestradiol (10−9 and 10−10 m), an increase of messenger levels of bFGF appears in hypothalamic type 1 astrocytes. Similarly, an induction of bFGF protein is also evident at this time of exposure. The effect on the mRNA and protein levels of bFGF is blocked by the presence in the medium of an antibody raised against the transforming growth factor α (TGFα) receptor. This observation indicates that, TGFα, whose synthesis is modulated by oestrogens in hypothalamic astrocytes and which is able to increase, both the mRNA and the protein levels of bFGF in our experimental model, may act as the mediator of the oestrogenic induction of bFGF. Hypothalamic astrocytes, together with hypothalamic neurones synthesizing and secreting luteinizing hormone‐releasing hormone (LHRH), form the LHRH network in conjunction with other neuronal systems. Gonadal steroids in general, and oestrogens in particular, play an important role in the control of the activity of this network. In addition, bFGF and TGFα, two growth factors released from astrocytes, are able to influence the activity of LHRH neurones. The present observations suggest that oestrogens may also act on LHRH neurones in an indirect fashion (i.e. by modulating the expression of bFGF and TGFα in glial cells).
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| [57] |
Gibberellic acid (GA) promotes seed germination, elongation growth, and flowering time in plants. GA responses are repressed by DELLA proteins, which contain an N-terminal DELLA domain essential for GA-dependent proteasomal degradation of DELLA repressors. Mutations of or within the DELLA domain of DELLA repressors have been described for species including Arabidopsis thaliana, wheat (Triticum aestivum), maize (Zea mays), and barley (Hordeum vulgare), and we show that these mutations confer GA insensitivity when introduced into the Arabidopsis GA INSENSITIVE (GAI) DELLA repressor. We also demonstrate that Arabidopsis mutants lacking the three GA INSENSITIVE DWARF1 (GID1) GA receptor genes are GA insensitive with respect to GA-promoted growth responses, GA-promoted DELLA repressor degradation, and GA-regulated gene expression. Our genetic interaction studies indicate that GAI and its close homolog REPRESSOR OF ga1-3 are the major growth repressors in a GA receptor mutant background. We further demonstrate that the GA insensitivity of the GAI DELLA domain mutants is explained in all cases by the inability of the mutant proteins to interact with the GID1A GA receptor. Since we found that the GAI DELLA domain alone can mediate GA-dependent GID1A interactions, we propose that the DELLA domain functions as a receiver domain for activated GA receptors.
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| [58] |
We investigated the physiological function of three Arabidopsis thaliana homologs of the gibberellin (GA) receptor GIBBERELLIN-INSENSITIVE DWARF1 (GID1) by determining the developmental consequences of GID1 inactivation in insertion mutants. Although single mutants developed normally, gid1a gid1c and gid1a gid1b displayed reduced stem height and lower male fertility, respectively, indicating some functional specificity. The triple mutant displayed a dwarf phenotype more severe than that of the extreme GA-deficient mutant ga1-3. Flower formation occurred in long days but was delayed, with severe defects in floral organ development. The triple mutant did not respond to applied GA. All three GID1 homologs were expressed in most tissues throughout development but differed in expression level. GA treatment reduced transcript abundance for all three GID1 genes, suggesting feedback regulation. The DELLA protein REPRESSOR OF ga1-3 (RGA) accumulated in the triple mutant, whose phenotype could be partially rescued by loss of RGA function. Yeast two-hybrid and in vitro pull-down assays confirmed that GA enhances the interaction between GID1 and DELLA proteins. In addition, the N-terminal sequence containing the DELLA domain is necessary for GID1 binding. Furthermore, yeast three-hybrid assays showed that the GA-GID1 complex promotes the interaction between RGA and the F-box protein SLY1, a component of the SCFSLY1 E3 ubiquitin ligase that targets the DELLA protein for degradation.
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| [59] |
<p>Brassinosteroids (BRs) are a group of naturally-occurring steroidal phytohormones playing fundamental roles during normal plant growth and development. Using a combination of experimental approaches, including analytical chemistry, genetics, and biochemistry, the major BR biosynthetic pathway has been largely elucidated. The least-understood knowledge in the BR research field is probably the molecular mechanisms controlling the bioactive levels of BRs in response to various developmental and environmental cues. In this review, we focus our discussion on a recently-proposed, 8-step predominant BR biosynthetic pathway, several newly-identified transcription factors regulating the expression of key enzymes that catalyze BR biosynthesis, and up-to-date information about the mechanisms that plants use to inactivate unnecessary BRs.</p><p><strong>Zhao B, Li J</strong> (2012) Regulation of brassinosteroid biosynthesis and inactivation. <em>J. Integr. Plant Biol</em>. <strong>54</strong>(10), 746–759.</p>
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Brassinosteroids (BRs) are plant hormones involved in various growth and developmental processes. The BR signaling system is well established in Arabidopsis (Arabidopsis thaliana) and rice (Oryza sativa) but poorly understood in maize (Zea mays). BRASSINOSTEROID INSENSITIVE1 (BRI1) is a BR receptor, and database searches and additional genomic sequencing identified five maize homologs including duplicate copies of BRI1 itself. RNA interference (RNAi) using the extracellular coding region of a maize zmbri1 complementary DNA knocked down the expression of all five homologs. Decreased response to exogenously applied brassinolide and altered BR marker gene expression demonstrate that zmbri1-RNAi transgenic lines have compromised BR signaling. zmbri1-RNAi plants showed dwarf stature due to shortened internodes, with upper internodes most strongly affected. Leaves of zmbri1-RNAi plants are dark green, upright, and twisted, with decreased auricle formation. Kinematic analysis showed that decreased cell division and cell elongation both contributed to the shortened leaves. A BRASSINOSTEROID INSENSITIVE1-ETHYL METHANESULFONATE-SUPPRESSOR1-yellow fluorescent protein (BES1-YFP) transgenic line was developed that showed BR-inducible BES1-YFP accumulation in the nucleus, which was decreased in zmbri1-RNAi. Expression of the BES1-YFP reporter was strong in the auricle region of developing leaves, suggesting that localized BR signaling is involved in promoting auricle development, consistent with the zmbri1-RNAi phenotype. The blade-sheath boundary disruption, shorter ligule, and disrupted auricle morphology of RNAi lines resemble KNOTTED1-LIKE HOMEOBOX (KNOX) mutants, consistent with a mechanistic connection between KNOX genes and BR signaling. © 2015 American Society of Plant Biologists. All Rights Reserved.
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| [62] |
\n Agriculturally advantageous reduction in plant height is usually achieved by blocking the action or production of gibberellins. Here, we describe a different dwarfing mechanism found in maize\n brachytic2\n (\n br2\n ) mutants characterized by compact lower stalk internodes. The height reduction in these plants results from the loss of a P-glycoprotein that modulates polar auxin transport in the maize stalk. The sorghum ortholog of\n br2\n is\n dwarf3\n (\n dw3\n ), an unstable mutant of long-standing commercial interest and concern. A direct duplication within the\n dw3\n gene is responsible for its mutant nature and also for its instability, because it facilitates unequal crossing-over at the locus.\n
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王晓雯, 王媛媛, 冯蓓祺, 等. 水稻矮化多蘖突变体mtd2/htd1-1的鉴定与图位克隆[J]. 西南大学学报(自然科学版), 2024, 46(2):2-12.
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唐兰, 吴元奇. 玉米矮秆基因及调控机理的研究进展[J]. 北方农业学报, 2023, 51(5):23-30.
株高是玉米理想株型的重要指标,与产量密切相关,影响着植物光合利用率、抗倒伏性、收获指数等。植物激素通过影响细胞的分裂和伸长来改变玉米的节间长度和数目,从而调节玉米高度达到矮化的效果。文章综述了近年来控制玉米株高的QTL/基因定位和克隆的研究进展以及激素(GA<sub>3</sub>、IAA、BR)调控下矮化突变体的形成,以期为玉米育种生产和应用提供参考。
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王文秀, 王磊. 玉米矮秆基因研究进展[J]. 生物技术通报, 2018, 34(11):22-26.
玉米已成为我国种植面积最大的农作物,提高玉米的种植密度是提高玉米产量的重要技术手段。玉米的株型改良成为玉米遗传育种研究的重点之一。合理株型和抗倒伏能力是影响玉米产量的重要农艺性状,也是衡量优良玉米品种的重要指标之一。通过玉米矮化基因的分离及研究,开发相应的分子标记,培育出矮化或半矮化玉米植株,从而改善玉米株型结构,增加种植密度,提高群体光合效能,提高玉米产量。主要从玉米矮化育种的概况、玉米矮化基因遗传特点、玉米矮化基因的表型以及激素对玉米矮化的影响等方面的研究进行了总结和概述,为玉米矮化基因的开发利用及分子机制的阐明提供新的信息。
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