Trova il tuo prossimo book preferito
Abbonati oggi e leggi gratis per 30 giorniInizia la tua prova gratuita di 30 giorniInformazioni sul libro
Sprouted Grains: Nutritional Value, Production, and Applications
Azioni libro
Inizia a leggere- Editore:
- Elsevier Science
- Pubblicato:
- Oct 11, 2018
- ISBN:
- 9780128115268
- Formato:
- Libro
Descrizione
Sprouted Grains: Nutritional Value, Production and Applications is a complete and comprehensive overview of sprouted grains, with coverage from grain to product. Sections includes discussions on the process of grain germination from both a genetic and physiological perspective, the nutrients and bioactive compounds present in spouted grains, and the equipment and technical innovation of use to manufacturers of sprouted grains and sprouted grain products. This book is essential reading for cereal science academics and postgraduate students interested in the subject of cereal processing, but is also ideal for industrial product developers in cereal companies.
This edited volume brings together the world’s leading researchers on sprouted grains.
Presents the nutrient and bioactive components of these healthy grains Provides extensive coverage of products developed from sprouted grains Includes contributions from an International team of both academic and industrial authors Covers the equipment and technology used in grain processingInformazioni sul libro
Sprouted Grains: Nutritional Value, Production, and Applications
Descrizione
Sprouted Grains: Nutritional Value, Production and Applications is a complete and comprehensive overview of sprouted grains, with coverage from grain to product. Sections includes discussions on the process of grain germination from both a genetic and physiological perspective, the nutrients and bioactive compounds present in spouted grains, and the equipment and technical innovation of use to manufacturers of sprouted grains and sprouted grain products. This book is essential reading for cereal science academics and postgraduate students interested in the subject of cereal processing, but is also ideal for industrial product developers in cereal companies.
This edited volume brings together the world’s leading researchers on sprouted grains.
Presents the nutrient and bioactive components of these healthy grains Provides extensive coverage of products developed from sprouted grains Includes contributions from an International team of both academic and industrial authors Covers the equipment and technology used in grain processing- Editore:
- Elsevier Science
- Pubblicato:
- Oct 11, 2018
- ISBN:
- 9780128115268
- Formato:
- Libro
Correlati a Sprouted Grains
Anteprima del libro
Sprouted Grains
China
Preface
The action of sprouting and fermenting grains prior to their use in foods and beverages has a long history of benefit to humankind. In the eras when safe, potable water was not readily available, sprouting of grains followed by fermentation as part of brewing processes provided access to safe beverages. In past millennia, humankind also learned that delicious foods could be produced using fermentation processes, such as traditional rice or wheat-based fermented foods or producing miso from soybeans. To many people who consume fermented grains as beverages and foods, the health and nutritional benefits were likely accepted or taken for granted as they consumed those products as part of their regular diets.
In recent years there has been vigorous renewal of interest in the properties of sprouted grains. With the advent of advanced analytical technologies and expansion of the knowledge base on nutrition, phytochemical composition, functionality and effects of enzyme activities, food scientists and technologists have endeavored to take advantage of the benefits of sprouting and fermentation of grains. They are producing new ingredients to be developed into new and exciting food products from both a fragrance, flavor, and nutrition perspective.
This compilation brings together the current state-of-the-art, science, and technology pertaining to sprouted grains. It describes the pathway of pre-sprouted seed to the sprouted seed and their respective properties, the production and properties of ingredients derived from the sprouted grain, and, finally, edible food products and their associated properties produced from the sprouted grain ingredients.
Our authors have graciously shared their extensive knowledge regarding sprouted grains, and for this we are most grateful. Please keep them in mind should you need further knowledge and expertise beyond that which is presented in this discourse.
The Editors
1
Molecular mechanisms of seed germination
Pham Anh Tuan, Menghan Sun, Tran-Nguyen Nguyen, Seokhoon Park and Belay T. Ayele, Department of Plant Science, University of Manitoba, Winnipeg, MB, Canada
Abstract
Seed germination is a complex physiological process that starts with the uptake of water by the quiescent dry seed and terminates with radicle protrusion through the seed covering layers; seed dormancy is an adaptive trait that blocks the germination of seeds under favorable environmental conditions. The need for rapid and uniform germination has led to the development of crop cultivars with a low level of seed dormancy, making them prone to precocious germination/sprouting when wet conditions occur prior to harvest. Given the significance of seed germination and dormancy traits in crop agriculture, understanding the molecular mechanisms underlying seed germination and dormancy has been the focus of seed science research. This chapter highlights progress made in the understanding of molecular aspects of seed germination and dormancy, with emphasis on regulation of the metabolic aspects of gibberellin and abscisic acid, the two plant hormones that act as major players in the control of germination and dormancy.
Keywords
Abscisic acid; cereals; dormancy; germination; gene expression; gibberellin; seed
Chapter Outline
1.1 Seed dormancy and germination 1
1.1.1 Seed dormancy 1
1.1.2 Physiological events associated with the release of seed dormancy 3
1.1.3 Seed germination and the associated physiological events 4
1.2 Hormonal regulation of dormancy and germination 5
1.2.1 Gibberellins promotes seed germination 6
1.2.2 Gibberellins regulate seed dormancy and germination in cereals 9
1.2.3 Abscisic acid and its metabolic pathway 10
1.2.4 Abscisic acid regulates dormancy induction in developing seeds of dicot and cereal crops 12
1.2.5 Abscisic acid regulates seed dormancy maintenance and germination in dicots 13
1.3 Conclusions and perspectives 16
References 17
1.1 Seed dormancy and germination
Most higher plants begin their life cycle as a seed, and seed germination is considered the first adaptive decision in their development. It is typically prevented for a specified period of time, even under apparently optimal environmental conditions, by a trait known as seed dormancy. In agricultural crops, a certain degree of dormancy is a desirable trait because it prevents precocious germination or preharvest sprouting of seeds. However, a high level of seed dormancy is undesirable as this results in nonuniform germination and seedling establishment, and incurs extra costs for seed storage until dormancy is lost.
1.1.1 Seed dormancy
Seed dormancy is defined as the inability of intact viable seeds to complete germination under favorable environmental conditions (Gao and Ayele, 2014). Seed dormancy can be classified as primary dormancy and secondary dormancy, based on when the induction of dormancy occurs. Primary dormancy is established during seed development due to endogenous factors and/or environmental conditions experienced by the mother plant (Gao and Ayele, 2014). Seeds that exhibit primary dormancy can be released from the state of dormancy by a number of treatments, which include after-ripening, defined as a period of dry storage of freshly harvested seeds at room temperature; and cold stratification, which refers to seed hydration/imbibition at low temperatures. When faced with conditions that are unfavorable to germination, nondormant seeds may also enter into the state of dormancy, and this type of dormancy is referred to as secondary dormancy (Kermode, 2005). Primary dormancy is further categorized into five different types: physiological dormancy, morphological dormancy, morphophysiological dormancy, physical dormancy, and combinational dormancy (physiological and physical dormancy) (Baskin and Baskin, 2004). Physiological dormancy is the most common type of dormancy present in many plant species, including weeds and crop species. It is further divided into nondeep, intermediate, and deep dormancy, based on the depth of dormancy. An excised embryo of deep physiologically dormant seed either cannot grow at all or will produce abnormal seedlings. In contrast to deep dormancy, excised embryos of nondeep physiologically dormant seeds can easily germinate and produce normal seedlings. Excised embryos of seeds that exhibit intermediate physiological dormancy can eventually germinate and grow. Morphological dormancy is caused by undifferentiated or underdeveloped embryo, however, germination can occur after further development of the embryo (Heather et al., 2010; Baskin and Baskin, 2004) while morphophysiological dormancy is characterized by an underdeveloped embryo that also exhibits physiological dormancy. Physical dormancy is caused by the seed coat or seed covering layers that are impermeable to water or gas, thereby blocking germination, while combinational dormancy is the type of dormancy induced by the physical and physiological components of the seed.
Based on the causes of dormancy, physiological dormancy is classified as embryo-induced dormancy and seed coat–induced dormancy (Hilhorst et al., 2010). Embryo-induced dormancy can be caused by a number of factors related to the embryo, including immaturity or underdevelopment of the embryo and/or synthesis of germination inhibitor compounds within the embryo. As a result, an excised embryo is unable to grow. Seed coat–induced dormancy is caused by the tissues surrounding the embryo, such as the endosperm and/or testa, that confer restraints against embryo growth, by interfering with the leaching of germination inhibitors from the embryo, and by restricting gaseous exchange and water uptake by the seed. In cereal grains, the coleorhiza tissue that covers the embryonic root may also prevent the emergence of the radicle, thereby causing dormancy (Barrero et al., 2009). Both the embryo and the covering appear to cause dormancy in many plant species and are considered common occurrences, irrespective of plant families and genera (Baskin and Baskin, 1998). In cereals, other external covering layers enclosing the caryopsis, consisting of lemma and palea, are reported to confer additional constraint for the germination of the embryo (Gualano and Benech-Arnold, 2009).
1.1.2 Physiological events associated with the release of seed dormancy
Seeds can be released from the state of dormancy by a number of treatments, including a period of dry storage at room temperature, which is referred to as after-ripening; cold and warm stratification; and treatment with smoke, light, and nitrate (Bailly, 2004; Bethke et al., 2006; Kucera et al., 2005). The duration of after-ripening that releases seeds from the state of dormancy varies with species, from a few weeks as long as several months (Bewley and Black, 1994). Seed storage environmental factors including moisture, temperature, and oxygen also determine the efficiency and thereby the duration of after-ripening necessary for inducing dormancy loss. The role of after-ripening in releasing seed dormancy is associated with changes in the physiological state of the seed, such as gene and protein expression, production of reactive oxygen species, oxidative modification of seed stored gene transcripts and proteins, and changes in the metabolism and signaling of plant hormones. Comparative transcriptomic analysis between dormant and after-ripened seeds of Arabidopsis revealed that both seed samples exhibit equally active but different gene expression programs. For example, genes related to assembly of the translation machinery are repressed in dormant seeds, while abscisic acid (ABA)-responsive elements are enriched in gene sets of dormant seed samples (Cadman et al., 2006). Furthermore, transcriptional activation of genes involved in cell wall modification and mobilization of seed storage reserves, such as lipids, proteins, and carbohydrates, is evident in after-ripened seed samples. Similarly, after ripening–induced dormancy release in wheat seeds is associated with transcriptional activation of genes involved in DNA replication, nitrogen metabolism, cytoplasmic membrane-bound vesicle formation, jasmonate biosynthesis and cell wall modification, and transcriptional repression of genes involved in protein synthesis/activity inhibitors during imbibition (Gao et al., 2012). In barley, after ripening–mediated alterations in the expression of genes involved in several biological processes have been reported, including those related to jasmonate responses, cell wall modification, nitrate and nitrite reduction and mRNA stability (Barrero et al., 2009).
Moreover, after-ripening causes the accumulation of reactive oxygen species in the embryo axes in species such as sunflower, leading to carbonylation of specific embryo proteins (Oracz et al., 2007). The release of seed dormancy by dry after-ripening in sunflower and wheat is also found to associate with oxidative modification of distinct seed-stored mRNAs (Bazin et al., 2011; Gao et al., 2013); the oxidized mRNAs in wheat seeds appeared to represent oxidative phosphorylation, ribosome biogenesis, nutrient reservoir, and α-amylase inhibitor activities (Gao et al., 2013). Furthermore, dormancy release in wheat seeds due to after-ripening leads to changes in the seed proteome, including those involved in proteolysis, cellular signaling, translation, and energy metabolism; and redox status of proteins including those related to carbohydrate, energy and amino acid metabolism, synthesis of secondary metabolites, genetic information processing, and antioxidative defenses (Bykova et al., 2011; Gao et al., 2013). However, the expression of some proteins, including those representing soluble and granule bound starch synthases, β-amylase, and elongation factor 1β, are shown to be co-regulated by imbibition in both dormant and nondormant seeds (Park et al., 2013), suggesting that the regulation of these proteins is not required for the acquisition of germination potential.
The role of after-ripening in breaking seed dormancy is also modulated by changes in the metabolism and signaling of plant hormones, mainly that of gibberellin (GA) and ABA (Chitnis et al., 2014; Liu et al., 2013; Shu et al., 2016), which is dicussed in more detail below.
1.1.3 Seed germination and the associated physiological events
The process of germination starts with seed imbibition/uptake of water by the dry seed and terminates with radicle penetration through the seed covering layers (Bewley, 1997; Weitbrecht et al., 2011). Generally, water uptake by dry seeds exhibits three phases (Bewley, 1997). Phase I is characterized by the initial rapid water uptake by a dry seed that causes seed swelling and change in shape (Bewley, 1997). Membrane structure is perturbed by the fast rehydration, leading to leakage of low molecular weight metabolites and cellular solutes out of the seed; however, the membrane structure is repaired after a short period of hydration. During this phase of water uptake, a number of physiological processes are initiated, including protein synthesis from existing mRNA and resumption of respiratory activities such as glycolytic and oxidative pentose phosphate respiratory pathways. The resumption of respiratory activities is characterized by a huge increase of oxygen consumption and release of carbon dioxide within minutes of imbibition (Bewley, 1997). Phase I of water uptake is also associated with repair of DNA, which is damaged during the desiccation phase of seed development, and this DNA repair involves DNA ligase that is activated following imbibition (Bewley, 1997; Weitbrecht et al., 2011). Repair of existing mitochondria is also initiated during phase I of seed imbibition. Since dry seeds contain a low amount of adenosine triphosphate (ATP), repair of the mitochondria during imbibition plays a role in the production of ATP required for the germination process.
Once the seed water uptake rate slows down and becomes stable, germinating seeds enter into phase II (Bewley, 1997; Weitbrecht et al., 2011). A number of events take place during this phase, including continued repair of existing DNA and mitochondria, synthesis of new mitochondria, and synthesis of proteins from newly synthesized gene transcripts. Furthermore, the initiation of embryo expansion and the weakening of seed covering layers occur during this phase (Bewley, 1997). Radicle protrusion through the testa/seed coat, which is defined as germination, marks the end of phase II and the beginning of phase III (postgermination stage). Phase III is mainly characterized by the mobilization of storage reserves deposited in storage organs, such as the endosperm in cereals; this triggers further increases in water uptake, succeeded by seedling growth (Nonogaki et al., 2010). Cell division, DNA synthesis and elongation of radicle cell also occur during the third phase of water uptake.
Transcriptomic and proteomic analyses of imbibing seeds have provided important insights into the physiological processes or molecular functions underlying seed germination. Transcriptomic analysis of barley seeds during germination reveals the transient upregulation of genes involved in cell wall synthesis Regulatory components such as transcription factors and signaling and posttranscriptional components are transiently upregulated during the early phase of germination, while the late germination phase is characterized by transcriptional activation of genes representing the histone families and many metabolic pathways, including amino acid metabolism, nucleotide metabolism, and those related to cell division and cell cycle (An and Lin, 2011). An and Lin also indicated that induction of genes associated with photosynthesis and reserve mobilization characterize the postgermination phase. In contrast, genes involved in stress-related pathways and seed storage proteins are repressed during the entire period of germination. Proteomic study of germinating barley seeds indicated that the expression of embryonic β-type proteasome, heat shock, desiccation stress, and ABA-induced proteins increases during the early period of imbibition, but their expression level decreases as imbibition continues (Bønsager et al., 2007). This suggests that programming of germination is initiated during the seed maturation phase of seed development. Redox-related proteins, such as ascorbate peroxidase, and enzymes involved in the degradation of storage reserves, such α-amylase, appeared to exhibit upregulation as imbibition proceeds further, while the expression of some proteins, such as those representing enzymes involved in glycolysis, is evident throughout the entire germination process, likely to provide the energy required to support the growing embryo.
Combined transcriptomic and metabolomics analysis of germinating rice seeds showed rapid increases in the level of some metabolites, including hexose phosphates, tricarboxylic acid cycle intermediates, and gamma-aminobutyric acid, followed by transcriptional activation of a large group of genes enriched in metabolic functional categories, and this appeared to trigger changes in metabolome, such as those related to carbohydrate, amino acid and cell wall metabolism (Howell et al., 2009). Proteomic analysis of germinating rice seeds also indicated the importance of proteins related to metabolism, specifically those involved in reserve mobilization and synthesis of precursors, and the regulation of redox homeostasis and gene expression for the germination of rice seeds (He et al., 2011).
Analysis of the transcriptome of germinating wheat seeds showed changes in the expression of genes involved in storage reserve mobilization, oxidative phosphorylation, tricarboxylic acid cycle, mitochondrial electron transport, and cell wall metabolism (Yu et al., 2014). Furthermore, the germination of wheat seed has been shown as associated with differential expression of proteins involved in storage, carbohydrate and cell metabolism, and gene transcription and translation (Dong et al., 2015).
1.2 Hormonal regulation of dormancy and germination
Plant hormones are involved in the regulation of the induction, maintenance and release of seed dormancy (Bentsink and Koornneef, 2008; Gao and Ayele, 2014); ABA and GA play major roles in this respect. ABA regulates dormancy induction and maintenance (Kermode, 2005), while GA promotes seed dormancy decay and germination. In addition to ABA and GA, other plant hormones such as ethylene, cytokinin, brassinosteroid, auxin, and jasmonates (JA) have been implicated in the regulation of seed dormancy and germination. The role of ethylene in seed dormancy and germination is associated with its crosstalk with GA and ABA (Arc et al., 2013; Beaudoin et al., 2000; Corbineau et al., 2014). For example, ethylene insensitive mutants (etr, ein2 and ein6) are hypersensitive to ABA and fail to germinate (Corbineau et al., 2014). The expression of the ethylene biosynthetic genes was found to be induced in germinating seeds of ABA-insensitive mutants, compared to that observed in wild type seeds (Linkies et al., 2009). Cytokinin also contributes to seed dormancy release via enhancing ethylene biosynthesis (Babiker et al., 1993; Kucera et al., 2005), and promotes seed germination through antagonizing the germination inhibitory effects of ABA (Guan et al., 2014; Wang et al., 2011). Brassinosteroid is the other plant hormone that can promote seed germination through antagonizing the effects of ABA. Previous studies with Arabidopsis showed that ABA strongly inhibits the germination of brassinosteroid-deficient mutant, det2-1, and brassinosteroid-signaling mutant, bri1-1 compared to that of the wild type (Steber and McCourt, 2001). Several reports have also suggested the potential role of auxin in the maintenance of seed dormancy. For example, exogenous auxin inhibits seed germination in wheat (Morris et al., 1988; Ramaih et al., 2003) and auxin-ABA interaction has been implicated in the inhibition of seed germination (Brady et al., 2003; Liu et al., 2007). JAs appear to have effects on seed dormancy and germination; however, different roles are observed in different plant species (Linkies and Leubner-Metzger, 2012). Both synergistic and antagonistic interactions between JA and ABA have been reported to occur during germination (Berger et al., 1996; Ellis and Turner, 2002; Staswick et al., 1992). For example, a synergistic effect of JA and ABA occurs during germination of Arabidopsis and Brassica napus seeds (Wilen et al., 1991; Ellis and Turner, 2002) where as JA appears to reduce the level of ABA and dormancy in wheat seeds (Jacobsen et al., 2013). This chapter discusses progress in the understanding of the regulations of GA and ABA metabolic pathways, and therefore seed dormancy and germination, with particular emphasis on cereals.
1.2.1 Gibberellins promotes seed germination
GAs are a group of diterpenoid compounds that influence many plant growth and developmental processes, including seed dormancy and germination (Yamaguchi, 2008). At least 136 naturally occurring GAs have been identified from plant, fungi, and bacteria (Hedden and Sponsel, 2015); however, only a few GAs are biologically active, including GA1, GA3, GA4, and GA7, while the rest are either precursors or catabolic products of the precursor or bioactive GAs (Yamaguchi, 2008). All GAs are synthesized from geranyl geranyl diphosphate (GGDP), which acts as a common precursor for all the diterpenoids, and their biosynthetic pathway can be divided into three stages (Fig. 1.1). The first stage involves the conversion of GGDP to ent-kaurene through a two-step cyclization reaction mediated by ent-copalyl diphosphate synthase (CPS) and ent-kaurene synthase (KS). The second stage of GA biosynthesis involves the synthesis of GA12 through stepwise oxidation of ent-kaurene by the actions of ent-kaurene oxidase (KO) and ent-kaurenoic acid oxidase (KAO). The conversion of GA12 to bioactive GAs, mainly GA1 and GA4, represents the third and final stage of GA biosynthesis involving 13-hydroxylation or non-13-hydroxylation reactions, respectively, and a series of subsequent oxidation reactions that are catalyzed by 2-oxoglutarate-dependent dioxygenases, GA 20-oxidases (GA20ox) and 3-oxidases (GA3ox).
Figure 1.1 Gibberellin metabolism pathway in plants. GGDP, geranylgeranyl diphosphate; CPS, ent-copalyl diphosphate synthase; KS, ent-kaurene synthase; KO, ent-kaurene oxidase; KAO, ent-kaurenoic acid oxidase; GA13ox, GA 13-oxidase; GA20ox, GA 20-oxidase; GA3ox, GA 3-oxidase; GA2ox, GA 2-oxidase.
The amount of bioactive GAs in plant tissues is also controlled by their inactivation. GA 2-oxidation is considered the major form of reaction that converts bioactive GAs and/or their precursors into inactive forms, and this reaction is catalyzed by GA 2-oxidases (GA2ox). Other GA inactivation reactions include epoxidation of the 16, 17-double bond of non-13-hydroxylated GAs, which is catalyzed by a member of the P450 class of enzymes (designated as ELONGATED UPPERMOST INTERNODE (EUI) in rice) (Zhu et al., 2006), and methylation of the C-6 carboxyl group of GAs, which is catalyzed by GA methyl transferases (GAMT) that use both bioactive GAs and their precursors as substrates to produce the corresponding methyl esters (Varbanova et al., 2007). GAs can also be inactivated through conjugation, most commonly with glucose.
The presence of an excess amount of GA in plants has negative effects, such as causing excessive growth of plants and precocious germination/vivipary of seeds (Coles et al., 1999; Huang et al., 1998; White and Rivin, 2000). Accordingly, it is important for plants to maintain optimal levels of bioactive GAs to ensure normal growth and development. Enzymes involved in the later part of the GA metabolic pathway, GA20ox, GA3ox, and GA2ox, play important roles in the regulation of bioactive GA level in plant tissues, and genes encoding these enzymes have been isolated from many plant species and form multigene families (Hedden and Phillips, 2000; Yamaguchi, 2008). Expression and mutational analysis of these regulatory genes have provided important insights into the elucidation of molecular mechanisms underlying the control of ABA and GA levels in seeds, and thereby seed dormancy and germination.
The role of GA in enhancing seed germination is well demonstrated. Seeds from GA-deficient mutants of Arabidopsis (ga1-3) and tomato (gib-1) due to defective CPS are unable to germinate without treatment with exogenous GA (Groot and Karssen, 1987; Karssen et al., 1989). GA has two roles in controlling germination in dicot species with endospermic seeds such as Arabidopsis. The first one is to enhance the weakening of the endosperm through inducing the expression of genes involved in cell wall hydrolysis such as β-mannanase, expansin, β-1,3-glucanase, chitinase, and activity of the corresponding enzymes (Chen and Bradford, 2000; Nonogaki et al., 2000; Wu et al., 2001). Given that GA-deficient embryos exhibit a reduced growth rate (Groot and Karssen, 1987), stimulating the growth potential of the embryo is believed to be the second role of GA in the regulation of seed germination (Debeaujon and Koornneef, 2000). Furthermore, imbibition of nondormant seeds is associated with increased production of bioactive GA (Ogawa et al., 2003), while seed treatment with chemical compounds that block GA biosynthesis, such as tetcyclacis and paclobutrazol, inhibits their germination (Karssen et al., 1989; Nambara et al., 1991). Analysis of the temporal and spatial expression patterns of GA biosynthesis genes during germination of Arabidopsis seeds showed that expression of the early GA biosynthesis gene CPS1, encoding an enzyme that catalyzes the first step of GGDP cyclization, is localized in the embryo provasculature while that of KO1, GA3ox1 and GA3ox2 occurs in the endodermis and cortex of embryo axis that undergoes expansion during germination (Yamaguchi et al., 2001). These results, along with reduced growth observed in GA-deficient embryos (Groot and Karssen, 1987), imply intercellular transport of GA intermediates for the synthesis of bioactive GA at or close to the sites of its action during germination. The induction in the expression of GA biosynthetic genes, including KO1, GA20ox3 and GA3ox1, as well as the low level expression of the GA inactivating GA2ox genes during seed imbibition, implies the significance of GA biosynthesis rather than GA inactivation in seed germination (Ogawa et al., 2003).
1.2.2 Gibberellins regulate seed dormancy and germination in cereals
The role of GAs in seed dormancy and germination of cereal seeds has also been well documented. Comparison between maturing seeds from dormant and less dormant lines of sorghum revealed that the embryos of the dormant seeds exhibit increased expression of GA2ox genes and lower bioactive GA level during their imbibition, while increases in the expression of two GA20ox genes and bioactive GA level were apparent in the embryos of the less dormant line (Rodríguez et al., 2012). Treatment of maize seeds with exogenous bioactive GA results in a higher germination rate, compared to those imbibed with water (Tian et al., 2014), while treatment with paclobutrazol, inhibitor of GA biosynthesis, leads to inhibition of seed germination (Song et al., 2011). Expression analysis of GA biosynthetic genes during germination in maize embryos showed temporally distinct expression patterns of the GA20ox gene family members, in which the peak expression of GA20ox1, GA20ox4 and GA20ox5 occurred at 32, 72, and 40 h after imbibition, respectively, while the expression of GA3ox1 was found to peak at 72 h after imbibition (Song et al., 2011). These results highlight that germination of maize seeds is accompanied by increased synthesis of bioactive GAs. The expressions of the GA biosynthetic GA20ox (GA20ox1, GA20ox2 and GA20ox3) and GA3ox2 genes of rice are induced during imbibition of nondormant seeds (Du et al., 2015; Kaneko et al., 2002). Loss-of-function mutation in the rice GA20ox2 gene has been consistently shown to reduce seed GA level and result in enhanced seed dormancy (Ye et al., 2015). A genome-wide association mapping in rice has also revealed GA2ox3 as one of the candidate genes controlling preharvest sprouting/germination (Magwa et al., 2016).
After-ripening, one of the dormancy breaking treatments, has been reported to modulate the dynamics of GA metabolism. Increased expression level of the GA biosynthetic gene GA20ox3, along with higher germination rate and percentage, have been observed in after-ripened than dormant rice seeds (Du et al., 2015). Rapid induction in the expression of GA3ox2 was also shown during imbibition of after-ripened barley seeds, while it expressed at low level in the corresponding dormant seeds (Gubler et al., 2008). The expression of GA-inactivating GA2ox3 gene was also shown to increase following imbibition of barley seeds, irrespective of dormancy status; however, its expression decreases to low level with continued imbibition in after-ripened seeds. Previous studies have shown that the expression of GA biosynthetic genes, including GA20ox1 and GA3ox2 genes, is induced in the scutella of germinating wheat seeds (Appleford et al., 2006), leading to the production of bioactive GA, GA1 (Lenton et al., 1994). Dormancy release in wheat seeds by after-ripening has been consistently shown as associated with higher expression level of GA20ox and GA3ox during imbibition (Liu et al., 2013). These results suggest the role of after-ripening in regulating GA level, and thereby dormancy loss and germination.
1.2.2.1 Environmental regulation of gibberellin metabolism and seed germination
Environmental factors such as light and temperature can alter GA metabolism and therefore seed germination. The role of red light in inducing germination of Arabidopsis and lettuce seeds was shown to be related with upregulation of GA biosynthesis genes, GA3ox1 and GA3ox2, in Arabidopsis, and GA3ox1 in lettuce (Toyomasu et al., 1993; Yamaguchi et al., 1998) and repression of GA catabolic gene, GA2ox2 (Nakaminami et al., 2003; Seo et al., 2006; Yamauchi et al., 2007). Cold stratification of Arabidopsis seeds also increases the expression of GA biosynthetic genes, GA20ox (GA20ox1 and GA20ox2) and GA3ox (GA3ox1), and bioactive GA content, resulting in dormancy loss and subsequent germination (Yamauchi et al., 2004), while high temperature suppresses the expression of GA biosynthetic GA20ox (GA20ox1-3) and GA3ox (GA3ox1 and GA3ox2) genes and seed GA level, leading to inhibition of germination (Toh et al.,
Recensioni
Recensioni
Cosa pensano gli utenti di Sprouted Grains
00 valutazioni / 0 recensioni