RESUMO
BACKGROUND: The core regulation of the abscisic acid (ABA) signalling pathway comprises the multigenic families PYL, PP2C, and SnRK2. In this work, we conducted a genome-wide study of the components of these families in Cucurbita pepo. RESULTS: The bioinformatic analysis of the C. pepo genome resulted in the identification of 19 CpPYL, 102 CpPP2C and 10 CpSnRK2 genes. The investigation of gene structure and protein motifs allowed to define 4 PYL, 13 PP2C and 3 SnRK2 subfamilies. RNA-seq analysis was used to determine the expression of these gene families in different plant organs, as well as to detect their differential gene expression during germination, and in response to ABA and cold stress in leaves. The specific tissue expression of some gene members indicated the relevant role of some ABA signalling genes in plant development. Moreover, their differential expression under ABA treatment or cold stress revealed those ABA signalling genes that responded to ABA, and those that were up- or down-regulated in response to cold stress. A reduced number of genes responded to both treatments. Specific PYL-PP2C-SnRK2 genes that had potential roles in germination were also detected, including those regulated early during the imbibition phase, those regulated later during the embryo extension and radicle emergence phase, and those induced or repressed during the whole germination process. CONCLUSIONS: The outcomes of this research open new research lines for agriculture and for assessing gene function in future studies.
Assuntos
Proteínas de Arabidopsis , Cucurbita , Ácido Abscísico/farmacologia , Ácido Abscísico/metabolismo , Cucurbita/genética , Cucurbita/metabolismo , Estudo de Associação Genômica Ampla , Plantas/genética , Resposta ao Choque Frio , Regulação da Expressão Gênica de Plantas , Proteínas de Arabidopsis/genéticaRESUMO
The sex determination process in cucurbits involves the control of stamen or carpel development during the specification of male or female flowers from a bisexual floral meristem, a function coordinated by ethylene. A gain-of-function mutation in the miR164-binding site of CpCUC2B, ortholog of the Arabidopsis transcription factor gene CUC2, not only produced ectopic floral meristems and organs, but also suppressed the development of carpels and promoted the development of stamens. The cuc2b mutation induced the transcription of CpCUC2B in the apical shoots of plants after female flowering but repressed other CUC genes regulated by miR164, suggesting a conserved functional redundancy of these genes in the development of squash flowers. The synergistic androecious phenotype of the double mutant between cuc2b and etr2b, an ethylene-insensitive mutation that enhances the production of male flowers, demonstrated that CpCUC2B arrests the development of carpels independently of ethylene and CpWIP1B. The transcriptional regulation of CpCUC1, CpCUC2, and ethylene genes in cuc2b and ethylene mutants also confirms this conclusion. However, the epistasis of cuc2b over aco1a, a mutation that suppresses stamen arrest in female flowers, and the down-regulation of CpACS27A in cuc2b female apical shoots, indicated that CpCUC2B promotes stamen development by suppressing the late ethylene production.
Assuntos
Arabidopsis , Cucurbita , Cucurbita/genética , Arabidopsis/genética , Etilenos , Flores , Fatores de Transcrição/metabolismo , Mutação , Regulação da Expressão Gênica de Plantas , MeristemaRESUMO
The crosstalk of phytohormones in the regulation of growth and development and the response of plants to environmental stresses is a cutting-edge research topic, especially in crop species. In this paper, we study the role and crosstalk between abscisic acid (ABA), ethylene (ET), and jasmonate (JA) in the control of germination and seedling growth in water or in standard nutrient solution and under salt stress (supplemented with 100-200 mM NaCl). The roles of ET and JA were studied using squash ET- and JA-deficient mutants aco1a and lox3a, respectively, while the crosstalk between ET, JA, and ABA was determined by comparing the expression of the key ABA, JA, and ET genes in wild-type (WT) and mutant genotypes under standard conditions and salt stress. Data showed that ET and JA are positive regulators of squash germination, a function that was found to be mediated by downregulating the ABA biosynthesis and signaling pathways. Under salt stress, aco1a germinated earlier than WT, while lox3a showed the same germination rate as WT, indicating that ET, but not JA, restricts squash germination under unfavorable salinity conditions, a function that was also mediated by upregulation of ABA. ET and JA were found to be negative regulators of plant growth during seedling establishment, although ET inhibits both the aerial part and the root, while JA inhibits only the root. Both aco1a and lox3a mutant roots showed increased tolerance to salt stress, a phenotype that was found to be mainly mediated by JA, although we cannot exclude that it is also mediated by ABA.
Assuntos
Ácido Abscísico , Cucurbita , Ciclopentanos , Etilenos , Regulação da Expressão Gênica de Plantas , Germinação , Oxilipinas , Reguladores de Crescimento de Plantas , Estresse Salino , Ciclopentanos/metabolismo , Germinação/efeitos dos fármacos , Etilenos/metabolismo , Ácido Abscísico/metabolismo , Oxilipinas/metabolismo , Cucurbita/crescimento & desenvolvimento , Cucurbita/genética , Cucurbita/metabolismo , Reguladores de Crescimento de Plantas/metabolismo , Plântula/crescimento & desenvolvimento , Plântula/metabolismo , Plântula/genética , Plântula/efeitos dos fármacos , Transdução de Sinais , Proteínas de Plantas/metabolismo , Proteínas de Plantas/genéticaRESUMO
Jasmonate (JA) has been found to be a relevant hormone in floral development in numerous species, but its function in cucurbit floral development and sex determination is unknown. Crosstalk between JA and ethylene (ET) in the differential regulation of male and female floral development was investigated by using the novel JA-deficient mutant lox3a, and the ET-deficient and -insensitive mutants, aco1a and etr2b, respectively, of Cucurbita pepo. The lox3a mutation suppresses male and female flower opening and induces the development of parthenocarpic fruit. A bulked-segregant analysis coupled with whole genome sequencing and fine mapping approach allowed the identification of lox3a mutation in CpLOX3A, a LIPOXYGENASE gene involved in JA biosynthesis. The reduced JA content and expression of JA-signalling genes in male and female flowers of lox3a, and the rescue of lox3a phenotype by external application of methyl jasmonate (MeJA), demonstrated that JA controls petal elongation and flower opening, as well as fruit abortion in the absence of fertilization. JA also rescued the phenotype of ET mutants aco1a and etr2b, which are both specifically defective in female flower opening and fruit abortion. ET, the sex determining hormone of cucurbits, is induced in female flowers towards anthesis, activating JA production and promoting the aperture of the female flower, and the abortion of the unfertilized ovary. Given the close association between flower closure and parthenocarpic fruit development, we propose that flower opening can act as a switch that triggers fruit set and development in fertilized ovaries, but may alternatively induce the abortion of the unfertilized ovary. Both ET and JA from mature and senescent petals can serve as remote signals that determine the alternative development of the ovary and fruit.
Assuntos
Cucurbita , Frutas/metabolismo , Proteínas de Plantas/genética , Proteínas de Plantas/metabolismo , Etilenos/metabolismo , Flores , Ciclopentanos/metabolismo , Oxilipinas/metabolismo , Hormônios/metabolismo , Regulação da Expressão Gênica de PlantasRESUMO
The enhanced salt tolerance of squash ethylene-insensitive mutants during germination and early stages of seedling development suggested that abscisic acid (ABA) could mediate this tolerance. To gain insight into the crosstalk between ethylene and ABA in seed germination, the germination rate and early seedling growth of wild type (WT) and ethylene-insensitive etr2b mutant were compared in seeds germinated under water and exogenous ABA treatment. The etr2b seeds germinated earlier than WT under both water and ABA, and the effect of ABA on radicle length and seedling growth of etr2b was lower than in WT, indicating that etr2b is also insensitive to ABA. The comparison of ABA and ethylene contents and ABA and ethylene gene expression profiles in WT and etr2b dry and imbibed seeds in either water, NaCl or ABA demonstrated a clear crosstalk between ethylene and ABA in germination. The expression profiles of ethylene genes in WT and etr2b indicated that the role of ethylene in seed germination does not appear to follow the canonical ethylene signaling pathway. Instead, etr2b reduces ABA content during formation of the seeds (dry seeds) and in response to seed imbibition and germination, which means diminished dormancy in the ethylene mutant. The etr2b mutation downregulated the expression of ABA biosynthesis and signaling genes during germination, demonstrating the positive role of ethylene receptor gene CpETR2B on seed germination and early seedling growth in squash is mediated by ABA. The reduced effect of exogenous ABA on ethylene production and ethylene gene expression in etr2b seeds suggests that this regulation is also dependent on ethylene.
Assuntos
Ácido Abscísico , Cucurbita , Ácido Abscísico/metabolismo , Germinação , Reguladores de Crescimento de Plantas/metabolismo , Etilenos/metabolismo , Plântula/metabolismo , Mutação , Sementes , Água/metabolismo , Regulação da Expressão Gênica de PlantasRESUMO
In the monoecious Cucurbita pepo, the transition to female flowering is the time at which the plant starts the production of female flowers after an initial male phase of development. Ethylene plays an essential role in this process since some ethylene deficient and ethylene-insensitive mutants are androecious and only produce male flowers. To gain insight into the molecular mechanisms regulating the specification and early development of female flowers, we have compared the transcriptomic changes occurring in the shoot apices of WT and androecious ethylene-insensitive etr1b mutant plants upon female flowering transition. There were 1160 female flowering-specific DEGs identified in WT plants upon female flowering, and 284 of them were found to be modulated by the ethylene-insensitive etr1b mutation. The function of these DEGs indicated that female flower specification depends on the adoption of a transcriptional program that includes previously identified sex-determining genes in the ethylene pathway, but also genes controlling the biosynthesis and signaling pathways of other phytohormones, and those encoding for many different transcription factors. The transcriptomic changes suggested that gibberellins play a negative role in female flowering, while ethylene, auxins, ABA and cytokinins are positive regulators. Transcription factors from 34 families, including NAC, ERF, bHLH, bZIP, MYB and C2H2/CH3, were found to be regulating female flowering in an ethylene-dependent or -independent manner. Our data open a new perspective of the molecular mechanisms that control the specification and development of female flowers in C. pepo.
Assuntos
Cucurbita , Humanos , Reguladores de Crescimento de Plantas/metabolismo , Etilenos/metabolismo , RNA-Seq , Flores , Fatores de Transcrição/genética , Fatores de Transcrição/metabolismo , Regulação da Expressão Gênica de PlantasRESUMO
Ethylene is the key regulator of sex determination in monoecious species of the family Cucurbitaceae. This hormone determines which individual floral meristems develop as female or male flowers and the female flowering transition. The sex determination genes discovered so far code for ethylene biosynthesis enzymes, but little is known about the importance of ethylene signaling components. In this paper we characterize two novel ethylene-insensitive mutations (etr1a-1 and etr1b) which block the female flowering transition of Cucurbita pepo; this makes plants produce male flowers indefinitely (androecy). Two missense mutations in the ethylene-binding domain of the ethylene receptors CpETR1A or CpETR1B were identified as the causal mutations of these phenotypes by using whole-genome resequencing. The distinctive phenotypes of single and double mutants for four etr mutations have demonstrated that the final level of ethylene insensitivity depends upon the strength and dosage of mutant alleles for at least three cooperating ETR genes, and that the level of ethylene insensitivity determines the final sex phenotype of the plant. The sex phenotype ranges from monoecy in ethylene-sensitive wild-type plants to androecy in the strongest ethylene-insensitive ones, via andromonoecy in partially ethylene-insensitive plants. The induction of female flowering transition was found to be associated with upregulation of CpACS11, CpACO2 and CpACS27, three ethylene biosynthesis genes required for female flower development. A model is proposed herein, integrating both ethylene biosynthesis and receptor genes into the genetic network which regulates sex determination in C. pepo.
Assuntos
Cucurbita/crescimento & desenvolvimento , Flores/crescimento & desenvolvimento , Proteínas de Plantas/genética , Receptores de Superfície Celular/genética , Cucurbita/genética , Etilenos/metabolismo , Flores/genética , Frutas/crescimento & desenvolvimento , Genes de Plantas/genética , Genes de Plantas/fisiologia , Mutação/genética , Reguladores de Crescimento de Plantas/metabolismo , Proteínas de Plantas/fisiologia , Receptores de Superfície Celular/fisiologiaRESUMO
High-throughput screening of an ethyl methanesulfonate-generated mutant collection of Cucurbita pepo using the ethylene triple-response test resulted in the identification of two semi-dominant ethylene-insensitive mutants: etr1a and etr2b. Both mutations altered sex determination mechanisms, promoting conversion of female into bisexual or hermaphrodite flowers, and monoecy into andromonoecy, thereby delaying the transition to female flowering and reducing the number of pistillate flowers per plant. The mutations also altered the growth rate and maturity of petals and carpels in pistillate flowers, lengthening the time required for flowers to reach anthesis, as well as stimulating the growth rate of ovaries and the parthenocarpic development of fruits. Whole-genome sequencing allowed identification of the causal mutation of the phenotypes as two missense mutations in the coding region of CpETR1A and CpETR2B, each one corresponding to one of the duplicates of ethylene receptor genes highly homologous to Arabidopsis ETR1 and ETR2. The phenotypes of homozygous and heterozygous single- and double-mutant plants indicated that the two ethylene receptors cooperate in the control of the ethylene response. The level of ethylene insensitivity, which was determined by the strength of each mutant allele and the dose of wild-type and mutant etr1a and etr2b alleles, correlated with the degree of phenotypic changes in the mutants.
Assuntos
Cucurbita/genética , Flores/crescimento & desenvolvimento , Regulação da Expressão Gênica de Plantas/fisiologia , Proteínas de Plantas/genética , Receptores de Superfície Celular/genética , Cucurbita/crescimento & desenvolvimento , Flores/genética , Proteínas de Plantas/metabolismo , Receptores de Superfície Celular/metabolismoRESUMO
Sex determination in Rumex acetosa, a dioecious plant with a complex XY1 Y2 sex chromosome system (females are XX and males are XY1 Y2 ), is not controlled by an active Y chromosome but depends on the ratio between the number of X chromosomes and autosomes. To gain insight into the molecular mechanisms of sex determination, we generated a subtracted cDNA library enriched in genes specifically or predominantly expressed in female floral buds in early stages of development, when sex determination mechanisms come into play. In the present paper, we report the molecular and functional characterization of FEM32, a gene encoding a protein that shares a common architecture with proteins in different plants, animals, bacteria and fungi of the aerolysin superfamily; many of these function as ß pore-forming toxins. The expression analysis, assessed by northern blot, RT-PCR and in situ hybridization, demonstrates that this gene is specifically expressed in flowers in both early and late stages of development, although its transcripts accumulate much more in female flowers than in male flowers. The ectopic expression of FEM32 under both the constitutive promoter 35S and the flower-specific promoter AP3 in transgenic tobacco showed no obvious alteration in vegetative development but was able to alter floral organ growth and pollen fertility. The 35S::FEM32 and AP3::FEM32 transgenic lines showed a reduction in stamen development and pollen viability, as well as a diminution in fruit set, fruit development and seed production. Compared with other floral organs, pistil development was, however, enhanced in plants overexpressing FEM32. According to these effects, it is likely that FEM32 functions in Rumex by arresting stamen and pollen development during female flower development. The aerolysin-like pore-forming proteins of eukaryotes are mainly involved in defence mechanisms against bacteria, fungi and insects and are also involved in apoptosis and programmed cell death (PCD), a mechanism that could explain the role of FEM32 in Rumex sex determination.
Assuntos
Toxinas Bacterianas/genética , Flores/genética , Nicotiana/genética , Infertilidade das Plantas/genética , Proteínas de Plantas/genética , Proteínas Citotóxicas Formadoras de Poros/genética , Rumex/genética , Sequência de Aminoácidos , Toxinas Bacterianas/classificação , Flores/crescimento & desenvolvimento , Frutas/genética , Frutas/crescimento & desenvolvimento , Perfilação da Expressão Gênica/métodos , Regulação da Expressão Gênica no Desenvolvimento , Regulação da Expressão Gênica de Plantas , Filogenia , Proteínas de Plantas/classificação , Plantas Geneticamente Modificadas , Pólen/genética , Pólen/crescimento & desenvolvimento , Proteínas Citotóxicas Formadoras de Poros/classificação , Rumex/crescimento & desenvolvimento , Sementes/genética , Sementes/crescimento & desenvolvimento , Homologia de Sequência de Aminoácidos , Nicotiana/crescimento & desenvolvimentoRESUMO
The Cucurbita genus (squashes, pumpkins and gourds) includes important domesticated species such as C. pepo, C. maxima and C. moschata. In this study, we present a high-quality draft of the zucchini (C. pepo) genome. The assembly has a size of 263 Mb, a scaffold N50 of 1.8 Mb and 34 240 gene models. It includes 92% of the conserved BUSCO core gene set, and it is estimated to cover 93.0% of the genome. The genome is organized in 20 pseudomolecules that represent 81.4% of the assembly, and it is integrated with a genetic map of 7718 SNPs. Despite the small genome size, three independent lines of evidence support that the C. pepo genome is the result of a whole-genome duplication: the topology of the gene family phylogenies, the karyotype organization and the distribution of 4DTv distances. Additionally, 40 transcriptomes of 12 species of the genus were assembled and analysed together with all the other published genomes of the Cucurbitaceae family. The duplication was detected in all the Cucurbita species analysed, including C. maxima and C. moschata, but not in the more distant cucurbits belonging to the Cucumis and Citrullus genera, and it is likely to have occurred 30 ± 4 Mya in the ancestral species that gave rise to the genus.
Assuntos
Evolução Biológica , Cucurbita/genética , Duplicação Gênica , Genoma de Planta , Transcriptoma , Cucurbita/metabolismoRESUMO
Immature, vegetable-like fruits are produced by crops of great economic importance, including cucumbers, zucchini, eggplants and bell peppers, among others. Because of their high respiration rates, associated with high rates of dehydration and metabolism, and their susceptibility to chilling injury (CI), vegetable fruits are highly perishable commodities, requiring particular storage conditions to avoid postharvest losses. This review focuses on the oxidative stress that affects the postharvest quality of vegetable fruits under chilling storage. We define the physiological and biochemical factors that are associated with the oxidative stress and the development of CI symptoms in these commodities, and discuss the different physical, chemical and biotechnological approaches that have been proposed to reduce oxidative stress while enhancing the chilling tolerance of vegetable fruits.
Assuntos
Biotecnologia/métodos , Estresse Oxidativo/fisiologia , Temperatura Baixa , Frutas/metabolismo , Verduras/metabolismoRESUMO
A number of Cucurbita pepo genotypes showing instable monoecy or partial andromonoecy, i.e. an incomplete conversion of female into bisexual flowers, have been detected. Given that in melon and cucumber andromonoecy is the result of reduction of ethylene production in female floral buds, caused by mutations in the ethylene biosynthesis genes CmACS7 and CsACS2; we have cloned and characterized two related C. pepo genes, CpACS27A and CpACS27B. The molecular structure of CpACS27A and its specific expression in the carpels of female flowers during earlier stages of flower development suggests that this gene is the Cucurbita ortholog of CmACS7 and CsACS2. CpACS27B is likely to be a paralogous pseudogene since it has not been found to be expressed in any of the analyzed tissues. CpACS27A was sequenced in Bolognese (Bog) and Vegetable Spaghetti (Veg), two monoecious inbred lines whose F2 was segregating for partial andromonoecy. The Bog allele of CpACS27A carried a missense mutation that resulted in a substitution of the conserved serine residue in position 176 by an alanine. Segregation analysis indicated that this mutant variant is necessary but not sufficient to confer the andromonoecious phenotype in squash. In concordance with its involvement in stamen arrest, a reduction in CpACS27A expression has been found in bisexual flower buds at earlier stages of development. This reduction in CpACS27A expression was concomitant with a downregulation of other ethylene biosynthesis and signaling genes during earlier and later stages of ovary development. The role of CpACS27A is discussed regarding the regulation of ethylene biosynthesis and signaling genes in the control of andromonoecy-associated traits, such as the delayed maturation of corolla and stigma as well as the parthenocarpic development of the fruit.
Assuntos
Cucurbita/genética , Cucurbita/metabolismo , Regulação da Expressão Gênica de Plantas/fisiologia , Proteínas de Plantas/metabolismo , Sequência de Aminoácidos , Clonagem Molecular , Etilenos/biossíntese , Flores/fisiologia , Genótipo , Dados de Sequência Molecular , Mutação , Filogenia , Proteínas de Plantas/genética , Reprodução/genética , Reprodução/fisiologia , Reação em Cadeia da Polimerase Via Transcriptase ReversaRESUMO
In monoecious species, female flowering constitutes the developmental process that determines the onset and production of fruit and is therefore closely related to crop yield. This article presents the identification and phenotypic and molecular characterization of myb62, an ethylmethane sulfonate loss-of-function mutation that completely blocks the female floral transition, converting all female flowers into male flowers. BSA-seq analysis coupled with WGS showed that myb62 corresponds to a C>T transition in the coding region of the gene CpMYB62, generating a premature stop codon and a truncated transcription factor without its N-terminal effector domain. The myb62 phenotype was partially rescued by exogenous ethylene application, indicating that the function of CpMYB62 is mediated by ethylene. Different evidence supports this conclusion: first, the reduced ethylene production of the mutant, and second, the male flower productive phenotype of the double mutant between myb62 and the ethylene-insensitive mutant etr2b, which demonstrated that myb62 is epistatic over etr2b. Furthermore, transcriptomic analysis of WT and myb62 apical shoots confirmed that CpMYB62 regulates master sex-determining genes, upregulating those encoding the ethylene biosynthesis enzymes CpACO2B and CpACS27A and those encoding for transcription factors that promote the development of carpels(CpCRC), but downregulating those involved in the arrest of carpels (CpWIP1), In the gene network controlling sex determination in cucurbits, CpMYB62 occupies the most upstream position, activating ethylene and other sex determining genes involved in female flower determination in Cucurbita pepo.
RESUMO
A Cucurbita pepo mutant with multiple defects in growth and development has been identified and characterized. The mutant dwfcp displayed a dwarf phenotype with dark green and shrinking leaves, shortened internodes and petioles, shorter but thicker roots and greater root biomass, and reduced fertility. The causal mutation of the phenotype was found to disrupt gene Cp4.1LG17g04540, the squash orthologue of the Arabidopsis brassinosteroid (BR) biosynthesis gene DWF5, encoding for 7-dehydrocholesterol reductase. A single nucleotide transition (G > A) causes a splicing defect in intron 6 that leads to a premature stop codon and a truncated CpDWF5 protein. The mutation co-segregated with the dwarf phenotype in a large BC1S1 segregating population. The reduced expression of CpDWF5 and brassinolide (BL) content in most mutant organs, and partial rescue of the mutant phenotype by exogenous application of BL, showed that the primary cause of the dwarfism in dwfcp is a BR deficiency. The results showed that in C. pepo, CpDWF5 is not only a positive growth regulator of different plant organs but also a negative regulator of salt tolerance. During germination and the early stages of seedling development, the dwarf mutant was less affected by salt stress than the wild type, concomitantly with a greater upregulation of genes associated with salt tolerance, including those involved in abscisic acid (ABA) biosynthesis, ABA and Ca2+ signaling, and those coding for cation exchangers and transporters.
RESUMO
Jasmonates are growth regulators that play a key role in flower development, fruit ripening, root growth, and plant defence. The study explores the coordination of floral organ maturation to ensure proper flower opening for pollination and fertilization. A new mutant (jar1b) was discovered, lacking petal elongation and flower opening but showing normal pistil and stamen development, leading to parthenocarpic fruit development. The mutation also enhanced the elongation of roots while reducing the formation of root hairs. BSA sequencing showed that jar1b is a missense mutation in the gene CpJAR1B, which encodes the enzyme that catalyzes the conjugation between JA and the amino acid isoleucine. The loss of function mutation in CpJAR1B produced a deficiency in biologically active (+) -7-iso-jasmonoyl-L-isoleucine (JA-Ile), which was not complemented by the paralogous gene CpJAR1A or any other redundant gene. Exogenous application of methyl jasmonate (MeJA) demonstrated that jar1b is partially insensitive to JA in both flowers and roots. Further experimentation involving the combination of JA-Ile deficient and ethylene-deficient, and ET insensitive mutations in double mutants revealed that CpJAR1B mediated ET action in female petal maturation and flower opening, but JA and ET have independent additive effects as negative regulators of the set and development of squash fruits. CpJAR1B also regulated the aperture of male flowers in an ethylene-independent manner. The root phenotype of jar1b and effects of external MeJA treatments indicated that CpJAR1B has a dual role in root development, inhibiting the elongation of primary and secondary roots, but promoting the formation of root hairs.
Assuntos
Cucurbita , Ciclopentanos , Flores , Frutas , Oxilipinas , Proteínas de Plantas , Ciclopentanos/farmacologia , Ciclopentanos/metabolismo , Oxilipinas/farmacologia , Oxilipinas/metabolismo , Flores/genética , Flores/crescimento & desenvolvimento , Flores/efeitos dos fármacos , Cucurbita/genética , Cucurbita/crescimento & desenvolvimento , Frutas/genética , Frutas/crescimento & desenvolvimento , Frutas/metabolismo , Proteínas de Plantas/genética , Proteínas de Plantas/metabolismo , Isoleucina/farmacologia , Isoleucina/metabolismo , Isoleucina/análogos & derivados , Mutação , Regulação da Expressão Gênica de Plantas/efeitos dos fármacos , Acetatos/farmacologia , Raízes de Plantas/crescimento & desenvolvimento , Raízes de Plantas/genética , Raízes de Plantas/metabolismo , Raízes de Plantas/efeitos dos fármacos , Reguladores de Crescimento de Plantas/metabolismo , Reguladores de Crescimento de Plantas/farmacologiaRESUMO
BACKGROUND: We have identified a kind of parthenocarpy in zucchini squash which is associated with an incomplete andromonoecy, i.e. a partial conversion of female into bisexual flowers. Given that andromonoecy in this and other cucurbit species is caused by a reduction of ethylene production in the female flower, the associated parthenocarpic development of the fruit suggested the involvement of ethylene in fruit set and early fruit development. RESULTS: We have compared the production of ethylene as well as the expression of 13 ethylene biosynthesis and signalling genes in pollinated and unpollinated ovaries/fruits of two cultivars, one of which is parthenocarpic (Cavili), while the other is non-parthenocarpic (Tosca). In the latter, unpollinated ovaries show an induction of ethylene biosynthesis and ethylene signal transduction pathway genes three days after anthesis, which is concomitant with the initiation of fruit abortion and senescence. Fruit set and early fruit development in pollinated flowers of both cultivars and unpollinated flowers of Cavili is coupled with low ethylene biosynthesis and signalling, which would also explain the partial andromonoecy in the parthenocarpic genotype. The reduction of ethylene production in the ovary cosegregates with parthenocarpy and partial andromonoecy in the selfing progeny of Cavili. Moreover, the induction of ethylene in anthesis (by ethephon treatments) reduced the percentage of bisexual parthenocarpic flowers in Cavili, while the inhibition of ethylene biosynthesis or response (by AVG and STS treatments) induces not only andromonoecy but also the parthenocarpic development of the fruit in both cultivars. CONCLUSIONS: Results demonstrate that a reduction of ethylene production or signalling in the zucchini flower is able to induce fruit set and early fruit development, and therefore that ethylene is actively involved in fruit set and early fruit development. Auxin and TIBA treatments, inducing fruit set and early fruit development in this species, also inhibit ethylene production and the expression of ethylene biosynthesis and response genes. A model is presented that discusses the crosstalk between ethylene and auxin in the control of fruit set and early fruit development in zucchini squash.
Assuntos
Cucurbita/metabolismo , Etilenos/biossíntese , Frutas/metabolismo , Cucurbita/crescimento & desenvolvimento , Cucurbita/fisiologia , Flores/efeitos dos fármacos , Flores/crescimento & desenvolvimento , Flores/metabolismo , Flores/fisiologia , Frutas/efeitos dos fármacos , Frutas/crescimento & desenvolvimento , Frutas/fisiologia , Regulação da Expressão Gênica de Plantas/efeitos dos fármacos , Ácidos Indolacéticos/farmacologia , Polinização/genética , Polinização/fisiologia , Ácidos Tri-Iodobenzoicos/farmacologiaRESUMO
The squash gain-of-function mutant etr2b disrupts the ethylene-binding domain of ethylene receptor CpETR2B, conferring partial ethylene insensitivity, changes in flower and fruit development, and enhanced salt tolerance. In this paper, we found that etr2b also confers a growth advantage as well as a physiological and metabolomic response that make the mutant better adapted to drought. Mutant plants had a higher root and leaf biomass than WT under both well-watered and drought conditions, but the reduction in growth parameters in response to drought was similar in WT and etr2b. Water deficit reduced all gas-exchange parameters in both WT and etr2b, but under moderate drought the mutant increased photosynthesis rate in comparison with control conditions, and showed a higher leaf CO2 concentration, transpiration rate, and stomata conductance than WT. The response of etr2b to drought indicates that ethylene is a negative regulator of plant growth under both control and drought. Since etr2b increased ABA content in well-watered plant, but prevented the induction of ABA production in response to drought, it is likely that the etr2b response under drought is not mediated by ABA. A 1H NMR metabolomic analysis revealed that etr2b enhances the accumulation of osmolytes (soluble sugars and trigonelline), unsaturated and polyunsaturated fatty acids, and phenolic compounds under drought, concomitantly with a reduction of malic- and fumaric-acid. The role of CpETR2B and ethylene in the regulation of these drought-protective metabolites is discussed.
Assuntos
Secas , Folhas de Planta , Folhas de Planta/metabolismo , Água/metabolismo , Etilenos/metabolismo , Estresse Fisiológico/fisiologia , Ácido Abscísico/metabolismoRESUMO
Reports show that phytohormone abscisic acid (ABA) is involved in reducing zucchini postharvest chilling injury. During the storage of harvested fruit at low temperatures, chilling injury symptoms were associated with cell damage through the production of reactive oxygen species. In this work, we have studied the importance of different non-enzymatic antioxidants on tolerance to cold stress in zucchini fruit treated with ABA. The application of ABA increases the antioxidant capacity of zucchini fruit during storage through the accumulation of ascorbate, carotenoids and polyphenolic compounds. The quantification of specific phenols was performed by UPLC/MS-MS, observing that exogenous ABA mainly activated the production of flavonoids. The rise in all these non-enzymatic antioxidants due to ABA correlates with a reduction in oxidative stress in treated fruit during cold stress. The results showed that the ABA mainly induces antioxidant metabolism during the first day of exposure to low temperatures, and this response is key to avoiding the occurrence of chilling injury. This work suggests an important protective role of non-enzymatic antioxidants and polyphenolic metabolism in the prevention of chilling injury in zucchini fruit.
RESUMO
Within the Cucurbitaceae family, most of its species develop unisexual female and male flowers, either on the same plant (monoecy) or on different plants (dioecy). As in other plant families, these two sex morphotypes have evolved from hermaphrodite species; however, many evolutionary events have occurred in cucurbits allowing easy conversion from dioecy to monoecy and vice versa. The variability in sex morphotypes is higher in the domesticated species of the family, which together with recent advances in genomics, make cucurbits an ideal model to study the genetic and molecular mechanisms that control sex determination in plants. Conventional studies demonstrated that ethylene was the master regulator of sex determination in cucurbits, although some cultivated species may respond differently to ethylene action. In this article, we survey the new advances in hormonal and genetic control of sex determination in cucurbit species, control which establishes the ethylene biosynthesis and signaling genes as being those that determine the floral meristem towards a male, female or hermaphrodite flower. The interactions between these genes are integrated into a model that explains the occurrence and distribution of unisexal and hermaphrodite flowers within the different sex morphotypes. We underline the significance of this scientific progress with regard to breeding programs for agronomically-important sex-associated traits.
Assuntos
Flores , Melhoramento Vegetal , Evolução Biológica , Etilenos , Feminino , Flores/genética , Masculino , FenótipoRESUMO
Cucurbitaceae is one of the most economically important plant families, and includes some worldwide cultivated species like cucumber, melons, and squashes, and some regionally cultivated and feral species that contribute to the human diet. For centuries, cucurbits have been appreciated because of their nutritional value and, in traditional medicine, because of their ability to alleviate certain ailments. Several studies have demonstrated the remarkable contents of valuable compounds in cucurbits, including antioxidants such as polyphenols, flavonoids, and carotenoids, but also tannins and terpenoids, which are abundant. This antioxidant power is beneficial for human health, but also in facing plant diseases and abiotic stresses. This review brings together data on the antioxidant properties of cucurbit species, addressing the genetic and pre- and postharvest factors that regulate the antioxidant content in different plant organs. Environmental conditions, management, storage, and pre- and postharvest treatments influencing the biosynthesis and activity of antioxidants, together with the biodiversity of this family, are determinant in improving the antioxidant potential of this group of species. Plant breeding, as well as the development of innovative biotechnological approaches, is also leading to new possibilities for exploiting cucurbits as functional products.