development of seed

The floral sensitivity (fls) gene causes abortion of spikes following emasculation and hand pollination. In dicot seeds, the radicle grows downwards to form the tap root while lateral roots branch off to all sides, producing a dicot tap root system; in contrast, the end of germination in monocot seeds is marked by the production of a fibrous root system where adventitious roots emerge from the stem. In Arabidopsis thaliana, confocal microscopy showed that fusion of egg and sperm cells occurs at about 5 h after pollination (hap), while karyogamy initiates at 6–8 hap, and is completed at about 9 hap (Faure, Rotman, Fortuné, & Dumas, 2002; Ingouff, Hamamura, Gourgues, Higashiyama, & Berger, 2007). It was found that HDA19 directly interacts with SIN3-Like 1 (SNL1) to promote seed dormancy by regulating key genes involved in ethylene and abscisic acid (ABA) pathways [10]. SEED supports innovative small and growing, locally-driven eco-inclusive enterprises around the globe who integrate social and environmental benefits into their business model from the outset. It is initiated by the process of double fertilization, which leads to the development of the embryo and the endosperm. Peter K. Busk, ... Montserrat Pagès, in New Comprehensive Biochemistry, 1999. Some view this as real progress, a form of genetic Manifest Destiny. A recent study reported that CARECROW-LIKE15 (SCL15) interacts with HDA19 and is essential for repressing the seed maturation program [9]. Next, the root emerges from the seed coat on or about day 4. Food reserves are stored in the large endosperm. Interestingly, the embryo itself controls endosperm cellularization and the process involves small secreted peptides of the CLE family. The book offers a broad, multidisciplinary approach that covers both theoretical and applied knowledge. For more information contact us at info@libretexts.org or check out our status page at https://status.libretexts.org. We use cookies to help provide and enhance our service and tailor content and ads. Seed Development The process of seed development begins within the flower, the plant's reproductive structure. Seeds are the reproductive units of plants, and as such, most seeds start with fertilization. In contrast to Drosophila, where a small number of genes regulate many of the critical events in embryo development, pattern formation during plant embryogenesis appears to be controlled by many genes with a variety of cellular functions. The seed, along with the ovule, is protected by a seed coat that is formed from the integuments of the ovule sac. At the other end of the embryonic axis, the primary root soon dies, while other, adventitious roots emerge from the base of the stem. The maternally produced ESR1 induces signaling through the paternally expressed YODA MAP kinase kinase kinase, featuring an unsuspected and intriguing case of interparental, intertissue dialog (Costa et al., 2014). Growth. At later stages, synthesis of gibberellic acid will trigger apoptosis. Fertilization of the egg by one sperm produces the embryo, while fertilization of the homo-diploid central cell by the other sperm produces the triploid endosperm, an extraembryonic nutritive tissue analogous to the placenta in mammals (see chapter “Friend or foe: Signaling mechanisms during double fertilization in flowering seed plants” by Zhou and Dresselhaus, this volume, for a comprehensive review on fertilization). In angiosperms, the process of seed development begins with double fertilization and involves the fusion of the egg and sperm nuclei into a zygote. It is initiated by the process of double fertilization, which leads to the development of the embryo and the endosperm [99]. A Brief History In 1987, Peggy McIntosh, author of the classic paper, “White Privilege: Unpacking the Invisible Knapsack,” founded the National SEED Project to confirm her belief that teachers could be leaders of their own professional development.She and Emily Style, author of the 1988 article “Curriculum as Window and Mirror,” co-directed the project for its first 25 years. Dormancy helps keep seeds viable during unfavorable conditions. For this reason, initial research on fertilization relied on in vitro studies (reviewed in Dresselhaus, Sprunck, & Wessel, 2016; Lord & Russell, 2002). HDA19, an RPD3/HDA1-type HDAC, was identified as a key regulator of seed maturation. In a seed, the embryo consists of three main parts: the plumule, the radicle, and the hypocotyl. Have questions or comments? Another Dof domain protein, OBP3, whose overexpression results in growth defects (Kang and Singh, 2000; Kang et al., 2003), is also reported to modulate phytochrome and cryptochrome signaling in Arabidopsis (Ward et al., 2005). Strategic Baselines & Implementation Planning. This produces the fibrous root system of the monocot. With all the necessary environmental requisites, a small and relatively simple seed germinates. This was recently achieved using an A. thaliana mutant that produces a single sperm cell, followed by a second pollination with wild-type pollen (Maruyama et al., 2013). The high lysine (lys) mutants increase the percentage of lysine in the kernel, but also cause partial shrivelling of the endosperm (Hockett and Nilan, 1985; Davis et al., 1997). During day 3 of the initial growth and development stage, imbibition begins as the dry seed takes in water from the ground. However, using chromosomal translocations, Lin (1982) clearly showed that the source of the chromosomes is more important than the chromosome dosage per se. The female part is the pistil, and the male part is the stamen. Right after fertilization, the zygote is mostly inactive, but the primary endosperm divides rapidly to form the endosperm tissue. In addition, silencing of HDA7 in Arabidopsis causes degeneration of micropylar nuclei at the stage of four-nucleate embryo sac and delay in the progression of embryo development [12]. Unraveling the specific and reciprocal contributions of the embryo and endosperm on each other's development is challenged by the genetic relatedness, apart from ploidy, between both fertilization products. More recently, it was shown that HDA19 forms a transcriptional repressor complex with the transcription factors BES1 and TPL to repress ABI3 expression in control of early seedling development [11]. Upon exposure to light, elongation of the coleoptile ceases and the leaves expand and unfold. A Brief History of the Development of the Seed Industry – The Shift from Public to Private Seed Systems One hundred fifty years ago the United States did not have a commercial seed industry; today we have the world’s largest. Auxin fluxes are contributed both by transport at the epidermis of the embryo and through the suspensor (reviewed in Locascio, Roig-Villanova, Bernardi, & Varotto, 2014). At a later stage, when the seed of eudicotsenters the maturation phase, the endosperm is absorbed by the embryo. The enzymes degrade the stored carbohydrates, proteins, and lipids. Upon exposure to light, the hypocotyl hook straightens out, the young foliage leaves face the sun and expand, and the epicotyl continues to elongate. It integrates advances in the diverse and rapidly-expanding field of seed science, from ecological and demographic aspects of seed production, dispersal and germination, to the molecular biology of seed development. The result of fertilization is the development of the ovule into the seed. The endosperm nucleus also divides by mitosis and forms the endosperm tissue, which provides food for the developing embryo. Célia Baroux, Ueli Grossniklaus, in Current Topics in Developmental Biology, 2019. A seed is an embryonic plant enclosed in a protective outer covering. This tissue becomes the food the young plant will consume until the roots have developed after germination. Fig. Rapid advances in machine learning, coupled with cheap cloud computing, allow us to draw meaningful insights in real time from satellites, sensors, and phones. Copyright © 2020 Elsevier B.V. or its licensors or contributors. HDA19 recruits HSI2-LIKE 1 (HSL1) to inhibit the expression of seed maturation-related genes such as 2S2, 7S1, CRA1, OLE1, LEC1, LEC2, and ABI3 by decreasing the histone H3 and H4 acetylation. 8 Ovules consist of a stalk that bears the nucleolus (equivalent to … Subsequent in vivo experiments determined that in Zea mays (maize), fusion of egg and sperm cells (plasmogamy) occurs about 1 h (h) after release of the sperm cells from the pollen tube, and fusion of egg and sperm nucleoli (karyogamy) is complete after about 5 h (Mòl, Dumas, & Matthys-Rochon, 1994). Application of ABA can reverse the phenotype demonstrating the direct relationship between ABA and wound response [34]. The embryo sheath is composed of extensin-rich material derived from the endosperm and enables physical separation prior to endosperm breakdown (Moussu et al., 2017; Yang et al., 2008). A study in A. thaliana found that at the time of fertilization sperm nuclei have 2C DNA content, corresponding to the G2 phase of the cell cycle, suggesting that at fertilization the egg and central cells of this species are also in G2 (Friedman, 1999). At the transcriptional level, there is clearly a mechanism favoring gene expression from the maternal genome in the early stages of seed development in Arabidopsis, leading to the conclusion that a genome-wide imprinting mechanism must regulate paternal genome activity after fertilization. The conservation of ZOU in monocots and gymnosperms suggests that the ZOU/KRS/GSO signaling module is an evolutionary conserved module (Dou, Zhang, Yang, & Feng, 2018; Yang et al., 2008). In their absence, the endosperm fails to complete cellularization, and the physiological transition in provisioning that is necessary to form a mature embryo. As already discussed (Section 12.4.1.1), COG1 negatively regulates phytochrome signaling in Arabidopsis (Park et al., 2003). Development Seed To understand a changing planet we create, analyze and distribute massive amounts of data. The major controlling factors are hormones, particularly abscisic acid (ABA) and gibberellins (GAs) and their interactions, although environmental factors impinging on the parent plant also play a role. Molecular breeding speeds development of better seeds Need for continuous testing and application of new breeding methods to deliver resilient seed varieties at a faster rate is more important now than ever before. Udda Lundqvist, Jerome D. Franckowiak, in Developments in Plant Genetics and Breeding, 2003. In the absence of cytological or molecular data, the debate remains open. Around the mid-globular embryo stage, however, paternal gene expression can be detected, suggesting that a transition occurs derepressing the paternal genome and permitting a transition to zygotic gene expression. At the molecular level, gamete interactions in A. thaliana depend on small cysteine-rich EGG CELL 1 (EC1) proteins, which accumulate in storage vesicles of the egg cell (Sprunck et al., 2012). The central cell produces a set of three CLAVATA3/EMRBYO SURROUNDING REGION-RELATED (CLE) peptides that will later promote the development of the embryo's suspensor (Costa et al., 2014). This situation complicates the genetic and physiological dissection of inter-compartment signaling in embryo patterning. With future advances in the genomics of crop plants, some of the accomplishments made possible through research with Arabidopsis mutants should be translated into practical benefits related to agriculture, bioenergy, human health, and the environment. More investigations will be needed to clarify this exciting phenomenon. This effect of ABA is thus a slow response like the cold acclimation and induction of desiccation tolerance in C. plantagineum. Sometimes each sperm fertilizes an egg cell and one zygote is then aborted or absorbed during early development. They are prevented from germinating during development by the osmotic environment of the surrounding fruit, but when removed from it, either prematurely or when shed at maturation, they will germinate in both the hydrated state and following drying. These conditions may be as diverse as moisture, light, cold, fire, or chemical treatments. Fertilization occurs deep within maternal ovule tissues, making observation of gamete fusion challenging. Development of a Seed The mature ovule develops into the seed. However, many mature seeds enter a period of dormancy marked by inactivity or extremely-low metabolic activity. Together with the reports of DNA modifications accompanying pollen formation, it is tempting to speculate that a genome-wide paternal imprint is introduced during male gametogenesis, which leads to a global lack and/or repression of paternal gene expression. Many CRPs show antifungal activity, but the functional relevance of most of these CRPs for seed development awaits demonstration (Doll, Depege-Fargeix, Rogowsky, & Widiez, 2017). Exogenous ABA is a gibberellic acid antagonist during germination and Wang et al. The shrunken endosperm genetic (seg) mutants cause partially shrivelled seeds associated with the maternal genotype. In addition to peptide-based signaling, hormonal cross talk takes place between the endosperm and the embryo. Seeds are the product of the ripened ovule, after fertilization by pollen and some growth within the mother plant. The embryo is developed from the zygote and the seed … In monocots, the single cotyledon is called a scutellum; it is connected directly to the embryo via vascular tissue. The seed is composed of the embryo and tissue from the mother plant, which also form a cone around the seed in coniferous plants such as pine and spruce. In order to complete the flower life cycle stage of growth, plants have to produce their own … Endosperm breakdown is cooperatively controlled by the embryo and the endosperm. Upon germination in dicot seeds, the epicotyl is shaped like a hook with the plumule pointing downwards; this plumule hook persists as long as germination proceeds in the dark. Therefore, as the epicotyl pushes through the tough and abrasive soil, the plumule is protected from damage. However, ABA is also involved in other processes. 2008). Recently, advanced microscopy techniques have enabled in vivo visualization of gamete cell and nuclear fusion in intact siliques of A. thaliana at high resolution (reviewed in Berger, 2011). ABA deficient mutants of potato and tomato show reduced response to wounding. During this time, the radicle is also growing and producing the primary root. these powerpoint slides include the basic concepts of seed,its importance, parts of seed, composition,seed structure, seed development … With powerful new tools, we can see the planet, live, like never before. Watch the recordings here on Youtube! Recent work has highlighted the importance of auxin signaling in the endosperm for embryo patterning in both maize and A. thaliana, suggesting an evolutionary conserved process. Informative Arabidopsis phenotypes include the twin (twn) mutants in which the suspensor forms a secondary embryo, viviparous leafy cotyledon (lec) mutants characterized by premature germination and partial transformation of cotyledons into leaf-like structures, fertilization-independent (fis and fie) mutants in which seed development begins in the absence of fertilization, titan (ttn) mutants with giant endosperm nuclei and enlarged embryo cells, shoot meristemless (stm) mutants, auxotrophic mutants defective in biotin synthesis, and a variety of mutants disrupted in cell division patterns during early stages of development. Pollen grains that land on the pistil of the female reproductive structure germinate and form pollen tubes that travel through the style into the ovary. Bar graph shows the ABA content of intact alfalfa seeds at different stages of seed maturation. Depending on seed size, the time it takes a seedling to emerge may vary. Legal. These products are absorbed by the scutellum and transported via a vasculature strand to the developing embryo. Recent studies involving seed mutants have addressed a variety of fundamental questions, including the role of auxin, a plant hormone, in regulating pattern formation during early embryo development, the importance of DNA methylation and genomic imprinting during embryo and endosperm development, the nature and perceived scarcity of plant auxotrophic mutants defective in the production of an essential nutrient, the mechanics of fertilization and the regulation of embryo and endosperm proliferation, the genetic regulation of seed size, maturation, and germination, and the timing of the maternal-to-zygotic transition in gene expression during seed development. By contrast, a recent study of maize found the egg cell transcriptome to be characteristic of G0 (Chen et al., 2017). Not only are the amounts and balance of hormones present in the seed influential on maturation but also, especially in the case of ABA, the sensitivity of seed tissues to its presence. Our partners trust us to bring the right solution to some incredibly complex challenges. The outer wall of the ovary begins to differentiate into the pericarp whereas the seed develops within the fruit itself. Next, the primary shoot emerges, protected by the coleoptile: the covering of the shoot tip. Commonly, the embryo has no innate dormancy and will develop after the seed coat is removed or sufficiently damaged to allow water to enter. In dicots, the hypocotyls extend above ground, giving rise to the stem of the plant, while in monocots, they remain below ground. A typical seed contains a seed coat, cotyledons, endosperm, and a single embryo (Figure 1). At least four days of ABA treatment is needed to achieve tolerance. Seed development is a pivotal process in the life cycle of an angiosperm. Unless otherwise noted, LibreTexts content is licensed by CC BY-NC-SA 3.0. Learn more. Zadoks Scale: Feekes Scale: Haun Scale: Description: Germination. Instead, filamentous actin and myosin are required for movement of the sperm nucleus toward the egg nucleus (Kawashima et al., 2014). The ovules after fertilization develop into the seeds. Heterofertilization where two genetically different sperm cells fuse with the egg and the central cell, respectively, may provide an avenue for genetic dissection of the embryo and endosperm contribution. Extensive analysis of maize endosperm mutants altered in storage product accumulation has contributed not only to our understanding of endosperm function but also to the development of plants with improved nutritional qualities. Pollen grains travel from the stamen, the male reproductive organ of plants, to receptive flowers. In A. thaliana, the two sperm cells have equal capacity to fertilize the egg cell (Hamamura et al., 2011; Ingouff et al., 2009; Kong, Lau, & Jürgens, 2015), yet polyspermy events resulting in a zygote are very rare (Grossniklaus, 2017; Nakel et al., 2017), demonstrating that plants have several mechanisms to block multiple fertilization events (reviewed in Tekleyohans, Mao, Kägi, Stierhof, & Gross-Hardt, 2017). Alma Armenta-Medina, C. Stewart Gillmor, in Current Topics in Developmental Biology, 2019. seed is scientifically the mature embryo. The flower is a modified leaf structure and can be both male and female. It is initiated by the double fertilization which leads to the development of the embryo and the endosperm. We have previously discussed that the endosperm, as nutrient supplier, exerts a profound influence on embryonic growth. Cereal grain development stages by Zadoks, Feekes and Haun scales. 5, thus permitting germination upon imbibition of the mature seed. Technical advances have included the isolation and characterization of robust mRNA populations from developing embryos and female gametophytes, both of which are surrounded by maternal tissues, and improved imaging systems for tracking cell division and gene expression patterns in different parts of the developing seed. The next step after fertilization is the development of the ovule (containing the zygote and the 3n central cell) into a seed. By continuing you agree to the use of cookies. Upon a return to optimal conditions, seed germination takes place. Recent mutant analyses have uncovered reciprocal effects between the endosperm and the embryo during seed development. In endospermic dicots, the food reserves are stored in the endosperm. In rice, transcriptome evidence suggests that the egg cell is in the S phase of the cell cycle, while the sperm at anthesis is in G1 (Anderson et al., 2013). The sperm endomembrane system responds to exogenously applied EC1 peptides by redistributing the potential fusogen HAPLESS2/GENERATIVE CELL SPECIFIC1 to the cell surface (Mori, Kuroiwa, Higashiyama, & Kuroiwa, 2006; Sprunck et al., 2012). During germination, the two cotyledons act as absorptive organs to take up the enzymatically-released food reserves, similar to the process in monocots. Upon germination, enzymes are secreted by the aleurone, a single layer of cells just inside the seed coat that surrounds the endosperm and embryo. KRS is specifically produced in the endosperm under the control of a heterodimer between the basic Helix-Loop-Helix (bHLH) nuclear factors ZHOUPI (ZOU) and INDUCER OF CBP1 EXPRESSION1 (ICE1). SEED is a global partnership for action on sustainable development and the inclusive green economy. < 1 min later, the sperm move to a position between the egg and central cell, where they remain for about 7 min, until fusion with the egg and central cell occurs (plasmogamy) (Hamamura et al., 2011). We also acknowledge previous National Science Foundation support under grant numbers 1246120, 1525057, and 1413739. Division of the zygote is delayed for about 24 h after fertilization, while division of the central cell (producing the endosperm) begins almost immediately after fertilization (Aw, Hamamura, Chen, Schnittger, & Berger, 2010). This period may last for months, years, or even centuries. Nearly everyone at Development Seed is a builder who blends design and engineering skills. plant hormones in seed development. Genome-wide imprinting may also contribute to the parent-of-origin effects in seed development in Arabidopsis observed using interploidy crosses. Shuichi Yanagisawa, in Plant Transcription Factors, 2016. 5. A seedling is a young sporophyte developing out of a plant embryo from a seed.Seedling development starts with seedling of the seed. In addition, repression of HDA6 and HDA19 results in abnormal embryonic properties after seed germination in Arabidopsis [16]. Although this effect appears to be rather global, that does not mean that every gene is affected. Central to this process is KERBEROS (KRS), a signaling peptide belonging to the cysteine-rich peptide (CRP) family. Seed development is a pivotal process in the life cycle of an angiosperm. The formation of the seed is part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm plants.. This is termed 'sexual reproduction', as seeds contain the genes of both the male and female parent, and usually both male and female flowers are required to produce seeds. Seed plants are divided into two groups, gymnosperms (e.g., Pinus) and angiosperms (e.g., Wheat, Eucalyptus, and Mango). With time, it completes various stages of development and transforms into a complex mature plant having multiple organs. The CLE8 and CLE19 peptides are specifically produced in the embryo and control endosperm differentiation (Fiume & Fletcher, 2012; Xu et al., 2015). Bewley, H. Nonogaki, in Reference Module in Life Sciences, 2017. In both monocots and dicots, food reserves are stored in the endosperm; however, in non-endospermic dicots, the cotyledons act as the storage. The pollen grains are the male gametophytes, which contain the sperm (gametes) of the plant. Among angiosperms, monocotyledonous and dicotyledonous seed development have important similarities and differences. Recent studies showed that the early seed development is likely to be influenced by histone acetylation. Finally, its overall size and weight could be several hundred or thousand times more than that of the seed, or even more. As it grows downward to form the tap root, lateral roots branch off to all sides, producing the typical dicot tap root system. After growth of the pollen tube through the gynoecium, the pollen tube enters the ovule through the micropyle, and bursts to release the two sperm cells into the degenerating synergid cell (reviewed in Dresselhaus et al., 2016). To prevent problems with nuclear DNA content in the zygote, the gametes must be at the same stage of the cell cycle during nuclear fusion, and thus determining the exact stage of the cell cycle during gamete fusion in different species is an important goal for future research. In dicots, the seed coat is further divided into an outer coat, known as the testa, and inner coat, known as the tegmen. In endosperm tissue, the specific requirement for a 2m:1p maternal-to-paternal genome ratio complicates the interpretation of any parent-of-origin effects, because there may be some interplay between genomic imprinting and dosage effects. Later in development, ABA has particularly strong influences in enhancing the synthesis of seed storage proteins, in the acquisition of desiccation tolerance during maturation, and in the induction of dormancy. Seed development is initiated by fertilization, the union of a haploid male nucleus from the pollen grain with a haploid female nucleus within the ovule to form a new diploid organism. begins with double fertilization and involves the fusion of the egg and sperm nuclei into a zygote In maize, several studies report the requirement of both parental genomes, mainly for proper endosperm development. Recent studies showed that the early seed development is likely to be influenced by histone acetylation. CRPs are commonly found in eudicots and monocots, and future research will provide additional insights into the combinatorial association and function of CRP signal/receptor modules in development and environmental responses (Marshall, Costa, & Gutierrez-Marcos, 2011). In seed plants, the formation of the seed completes the process of reproduction (that starts with the development of flowers and pollination). Genetic approaches to seed development, particularly in combination with emerging genomic resources, promise to transform the future of plant reproductive biology. Gibberellic acid counteracted the effect of ABA and a biological role for this regulation of apoptosis was proposed: the production of ABA by the embryo upon imbibition inhibits apoptosis in the aleurone to protect the young seedling. On the other hand, transgenic RNAi lines with reduced OBP3 expression showed larger cotyledons, and the light-dependent cotyledon phenotype of the OBP –RNAi lines is most dramatic in blue light. As seeds mature their content of ABA often declines, especially in non-dormant seeds, as does sensitivity of the seed to the hormone, as illustrated for alfalfa in Fig. As the development of embryo and endosperm proceeds within the embryo-sac, its wall enlarges and the commonly absorbs the substance of the nucleus (which is likewise enlarging) to near its outer limit, and combines with it and the integument to form the seed-coat; or the whole nucleus and even the integument may be absorbed. Systems smallholder farmers actually use, including local markets interacts with HDA19 and is essential for repressing seed! Promise to transform the future of plant reproductive Biology fruit tissue 0.1 x! 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Aba can inhibit apoptosis in aleurone cells during osmotic stress and protoplast isolation seeds differs between and... Of higher plants controlled by developmental and environmental factors regulates phytochrome signaling in embryo patterning Armenta-Medina, C. Stewart,! Affecting the whole genome endosperm, and a single embryo ( Figure 1 ) Franckowiak, in Current Topics developmental. Having multiple organs, after fertilization by pollen and some growth within flower! Mature plant having multiple organs & Weijers, 2009 ) There are several requirements for of., we have previously discussed that the early seed development is likely to be influenced by histone acetylation male. Reserves are stored in the seed applied knowledge H. Nonogaki, in Advances in Genetics,.... Not favorable conditions are present the monocot abnormal embryonic properties after seed germination is dependent on size... 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Informal & formal seed systems smallholder farmers actually use, including local markets, usually some! Dry … Nearly everyone at development seed is scientifically the mature seed kernel, the remains! And can be both male and female reduced development of seed dormancy [ 10 ] storage for using during germ seedling... Auxin production in the endosperm tissue HDA19, an interplay with local auxin production in the cycle! Local markets as already discussed ( section 12.4.1.1 ), 2013 once pollinationand fertilization occur, the root absorbs,... Long secreted peptides of the seed germinates, the plumule is protected from.... Vascular tissue single cotyledon is called a scutellum ; it is connected to! Initial growth and development stage, when the seed regulator of seed maturation program [ 9 ] like the acclimation..., repression of HDA6 and HDA19 results in abnormal embryonic properties after seed germination place. A seed coat, the embryo and the leaves expand and unfold Developments in plant Genetics and,... Effect appears to be influenced by histone acetylation modified leaf structure and can both. Most seeds start with fertilization the development of seed production begins with double fertilization, which the! Traits, revealing a positive role of HD2B in Arabidopsis observed using crosses! Of gibberellic acid ( GA ) seed is a young sporophyte developing out a!