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Endosperm

The endosperm is a triploid nutritive tissue unique to the seeds of angiosperms (flowering plants), formed during double fertilization and serving as the primary source of stored nutrients—such as starch, proteins, and lipids—for the developing embryo and early seedling growth.^[1] This tissue surrounds the embryo within the seed, acting analogously to the yolk in animal eggs by providing essential energy and building materials until the seedling can perform photosynthesis.^[1] Unlike gymnosperms, where nutritive tissue derives solely from maternal gametophyte, the endosperm's biparental origin (with a 2:1 maternal-to-paternal genome ratio in most cases) confers evolutionary advantages, including hybrid vigor and balanced resource allocation between parental genomes.^[2] The formation of the endosperm occurs through double fertilization, a hallmark reproductive innovation of angiosperms, in which one sperm nucleus from the pollen tube fuses with the egg cell to produce the diploid (2n) zygote that develops into the embryo, while the second sperm nucleus fuses with the two haploid polar nuclei in the central cell of the female gametophyte (embryo sac), yielding the triploid (3n) endosperm precursor.^[1] This process ensures synchronous development of both embryo and endosperm, with the endosperm initiating as a syncytium (multiple nuclei in shared cytoplasm) before cellularizing into a storage tissue.^[2] Ploidy levels can vary evolutionarily; for instance, diploid (2n) endosperm occurs in basal angiosperms like water lilies (Nymphaeaceae), suggesting an ancestral state, while triploidy predominates in derived lineages.^[2] Endosperm development involves rapid proliferation and accumulation of reserves post-fertilization, often comprising up to 80% of the seed's mass in economically important crops like maize and wheat, where it forms the starchy kernel.^[2] Its primary function is to nourish the embryo during seed maturation and support germination by mobilizing stored nutrients, though in many species—such as beans and peas—the endosperm is largely absorbed by expanding cotyledons before maturity, leaving the seed non-endospermic.^[3] In contrast, persistent endosperm in cereals and palms sustains the seedling longer, highlighting adaptive diversity.^[3] Beyond nutrition, the endosperm influences seed viability, parental conflict over resource provisioning, and postzygotic isolation in hybrids, underscoring its role in angiosperm reproductive success and diversification.^[2]

Formation and Development

Double Fertilization

Double fertilization is a hallmark reproductive process unique to angiosperms, the flowering plants, in which a single pollen tube delivers two sperm cells that participate in two distinct fusion events within the female gametophyte. This mechanism ensures the coordinated development of both the embryo and the nutrient-rich endosperm tissue essential for seed viability. The process was first discovered by Sergei Nawaschin in 1898 while studying Lilium martagon and Fritillaria tenella, with independent confirmation by Léon Guignard in 1899 using similar species.^[4]^[5] The process begins when the pollen tube, guided by chemical signals from the female tissues, reaches the micropyle of the ovule and discharges its contents into one of the two synergid cells flanking the egg apparatus in the embryo sac. One sperm cell then fuses with the haploid egg cell (n) to form the diploid zygote (2n), which will develop into the embryo. Simultaneously or shortly thereafter, the second sperm cell migrates to the central cell, where it fuses with the two polar nuclei (typically 2n, though varying by species) to produce the triploid primary endosperm nucleus (3n). This double fusion event, known as syngamy for the zygote and triple fusion for the endosperm, distinguishes angiosperm reproduction from that of other seed plants.^[6]^[7] Synergid cells play a critical role in facilitating and regulating double fertilization. These accessory cells secrete peptide signals, such as LURE proteins, that attract the pollen tube to the ovule and promote its rupture for sperm release.^[8] Following pollen tube arrival, one synergid degenerates, creating a receptive space for the sperm cells while preventing additional pollen tubes from entering, thus blocking polyspermy and ensuring monospermy at both the egg and central cell. This degeneration is triggered by fertilization signals and involves programmed cell death pathways.^[9] The triploid genome of the endosperm introduces a 2:1 maternal-to-paternal allele ratio, which underpins genomic imprinting—a parent-of-origin-specific epigenetic regulation of gene expression. In this system, certain paternal alleles promote endosperm growth, while maternal alleles often repress it, balancing nutrient allocation and preventing overproliferation that could compromise seed development. This imprinting is mediated by DNA methylation and histone modifications, with disruptions leading to seed abortion in interspecies crosses.^[10]^[11] A typical diagram of double fertilization illustrates the embryo sac within the ovule, showing the pollen tube entering through the micropyle and penetrating a synergid cell. The two sperm nuclei are depicted as arrows: one fusing with the egg cell at the chalazal end to form the zygote, and the other combining with the central cell's polar nuclei to yield the primary endosperm nucleus. Surrounding structures, including the persistent synergid and antipodal cells, highlight the spatial organization, with labels indicating ploidy levels and fusion outcomes.^[6]

Endosperm Development

Following double fertilization, the endosperm initiates development as a triploid tissue derived from the fusion of two maternal polar nuclei and one sperm nucleus, entering an initial coenocytic phase characterized by rapid free nuclear divisions without cytokinesis, resulting in a shared multinucleate cytoplasm (coenocyte).^[12] This phase allows for expansive growth, with nuclei migrating and positioning within the expanding cytoplasm, often forming distinct domains such as micropylar and chalazal regions. In many angiosperms, this coenocytic stage persists for several divisions, enabling nutrient uptake and synchronization before structural organization. For instance, in cereals like maize (Zea mays), typically 128–512 free nuclei form during this period.^[13] Endosperm development progresses through distinct patterns of cellularization, classified into three main types based on the timing and mode of cell wall formation. In nuclear endosperm development, found in ~83% of dicot families such as Brassicaceae (Arabidopsis thaliana) and Euphorbiaceae (Croton spp.), the coenocytic phase dominates initially, with cell walls forming later from the periphery inward or micropylar to chalazal end, converting the coenocyte into a cellular tissue.^[14] Cellular endosperm development, observed in families like Nymphaeaceae and Annonaceae, involves synchronous cytokinesis accompanying each mitosis from the first division, yielding a fully cellular structure early on without a prolonged coenocytic stage.^[15] Helobial endosperm, an intermediate type found in about 34 families primarily among monocots such as Hydrocharitaceae and Alismatales (e.g., Limnocharis flava), features an initial cytokinesis of the first zygote cell into micropylar and chalazal chambers; the micropylar chamber then undergoes free nuclear divisions, while the chalazal chamber cellularizes immediately or remains uninucleate.^[15] These patterns influence overall endosperm architecture, with nuclear types often leading to larger, more uniform storage tissues.^[12] Several factors regulate endosperm development, including hormonal signals, nutrient dynamics, and programmed cell death (PCD). Auxins and cytokinins, modulated by epigenetic mechanisms like genomic imprinting, guide nuclear positioning, cellularization waves, and domain specification during the coenocytic phase.^[15] Nutrient allocation from maternal tissues via the seed coat and funiculus supports rapid growth, with transfer cells at the micropylar end facilitating solute uptake through wall ingrowths.^[12] PCD occurs in peripheral layers, such as embryo-surrounding cells and parts of the starchy endosperm, to optimize resource partitioning and prevent overproliferation, often triggered after endoreduplication cycles that amplify gene expression for storage synthesis.^[12] During maturation, the endosperm accumulates storage reserves tailored to seed type, with starchy endosperm in cereals like rice (Oryza sativa) and maize depositing starches, proteins (e.g., prolamins), and lipids in central regions, while aleurone layers differentiate peripherally to enclose these reserves.^[12] In orthodox seeds, the tissue undergoes desiccation, reducing water content to 5–15% for dormancy, accompanied by PCD in storage cells to mobilize reserves post-germination; however, in recalcitrant seeds, the endosperm remains hydrated. Abnormalities frequently arise in interspecific or interploidy hybrids due to genomic imprinting conflicts, where imbalances in parental gene expression disrupt cellularization or nutrient provisioning, leading to endosperm breakdown and seed abortion—e.g., in crosses between Arabidopsis thaliana and Arabidopsis arenosa, requiring embryo rescue techniques.^[16] Such barriers highlight the endosperm's sensitivity to ploidy ratios, with paternally imprinted genes often promoting overproliferation in incompatible unions.^[17]

Structure and Types

Cellular Types

The endosperm in angiosperms exhibits structural diversity based on the patterns of nuclear division and cell wall formation following double fertilization, resulting in distinct cellular organizations in mature seeds. These variations influence nutrient storage and embryo nourishment, with three primary types recognized: nuclear, cellular, and helobial. A rarer variant involves pronounced vacuolation in certain families.^[15] Nuclear endosperm, the most widespread type, is characterized by repeated mitotic divisions of the primary endosperm nucleus without immediate cell wall formation, producing a multinucleate coenocyte or syncytium that later undergoes peripheral cellularization to form a uniform tissue layer. This process allows for rapid proliferation and expansive growth before structural barriers develop, often resulting in a central vacuole surrounded by free nuclei. It predominates in many eudicots, such as Arabidopsis thaliana in the Brassicaceae family, and is also common in advanced monocots like those in the Poales; distribution spans approximately 156 dicotyledonous and 30 monocotyledonous families across angiosperm clades. Functionally, this type enhances storage efficiency through initial free-nuclear expansion, facilitating nutrient uptake via associated haustoria, though it may limit compartmentalization compared to walled structures, potentially affecting uniform embryo access during germination.^[15]^[18] In contrast, cellular endosperm features synchronous nuclear divisions accompanied by cytokinesis from the outset, yielding a brick-like array of walled cells that maintain structural integrity throughout development. This early wall formation produces a highly organized tissue, often with uniform cell files radiating from the center, as seen in some basal angiosperms such as Nymphaea in Nymphaeaceae. It occurs in about 135 dicotyledonous families, reflecting a distribution biased toward dicots and certain primitive lineages. The compartmentalized structure supports efficient reserve accumulation in discrete cells, improving embryo access by enabling targeted degradation and nutrient mobilization, which is advantageous for seeds requiring precise storage partitioning.^[15]^[18] Helobial endosperm represents an intermediate form, where the first division of the primary endosperm nucleus produces a wall that separates two chambers: the larger micropylar chamber undergoes free nuclear divisions akin to nuclear endosperm, while the smaller chalazal chamber develops cellularly and often functions as a haustorium. This differential organization leads to asymmetric tissue development, with the micropylar region storing reserves and the chalazal aiding absorption. It is primarily found in 34 monocotyledonous families, including primitive groups like Hydrocharitaceae (e.g., Elodea) and Aponogetonaceae, with limited occurrence in dicots such as Acanthaceae. Functionally, this type balances rapid nuclear growth with early compartmentalization, optimizing nutrient transfer to the embryo through specialized haustorial invasion, though it may constrain overall storage volume in smaller seeds.^[15] Rare variants include highly vacuolate endosperm, observed in families like Papaveraceae (e.g., Papaver rhoeas), where nuclear or cellular development culminates in extensive vacuolation, forming a watery, expansive tissue with large central vacuoles that enhance hydration and storage capacity before cellular maturation. This occurs sporadically across angiosperm clades, such as in certain basal eudicots, and correlates with improved embryo support in oily or farinaceous seeds by facilitating osmotic nutrient release, though it risks structural instability if degradation is untimely. Overall, these cellular types' distributions align with phylogenetic patterns, with nuclear prevailing in derived lineages and cellular/helobial in basal ones, influencing seed viability through tailored reserve organization.^[19]^[15]^[18]

Composition and Ploidy

The endosperm in most angiosperms is triploid (3n), arising from the fusion of two polar nuclei from the maternal central cell (contributing two maternal genomes, 2n) with a single sperm nucleus (contributing one paternal genome, n) during double fertilization.^[20] This 2m:1p genomic ratio ensures a balanced genetic constitution that supports nutrient storage and embryo nourishment. Ploidy levels can vary across angiosperm lineages; for instance, diploid (2n) endosperm occurs in species with a single maternal contribution in the central cell, such as those in the Oenothera-type embryo sac found in various angiosperm lineages, including some eudicots like Oenothera.^[11] In apomictic plants, where asexual seed formation predominates, endosperm ploidy may be reduced to 2n due to unfertilized diploid eggs and secondary nuclei in diplosporic or aposporic pathways.^[11] Polyploid species exhibit elevated ploidy, such as 4n or higher in tetraploids, reflecting the multiplied gametic contributions (e.g., 2m:2p in some cases), which can influence seed size and viability.^[11] The biochemical composition of endosperm is dominated by storage reserves tailored to species-specific needs, primarily carbohydrates, proteins, lipids, and minerals. Storage carbohydrates, mainly starch composed of amylose (linear α-1,4-glucan chains) and amylopectin (branched chains with α-1,6 linkages), constitute up to 70% of dry weight in cereal endosperms like wheat (63-72%) and sorghum (82-87%), serving as the primary energy reserve.^[21] Proteins, including prolamins (e.g., zeins in maize, gliadins in wheat) and globulins, form 10-15% of the composition and are stored in protein bodies within the starchy endosperm, providing amino acids for early seedling growth.^[22] In oilseed species such as castor (Ricinus communis), lipids accumulate prominently in the endosperm (up to 50-60% as triacylglycerols in oil bodies), replacing starch as the main reserve for energy mobilization.^[21] Minerals, particularly phosphorus stored as phytic acid in phytin bodies within aleurone cells, account for 1-2% and facilitate nutrient sequestration, though they can reduce bioavailability in mature seeds.^[21] Specialized structures within the endosperm enhance its storage and functional roles, particularly in monocots. The aleurone layer, an outer sheath of one to three cuboidal, protein-rich cells (containing ~50% dietary fiber in wheat, along with lipids, vitamins, and minerals like iron and zinc), surrounds the inner starchy endosperm core and remains viable post-maturity.^[23] During germination, aleurone cells secrete hydrolytic enzymes such as α-amylase and lipase, activated by gibberellin from the embryo, to degrade reserves in the adjacent starchy endosperm, which consists of dead cells packed with starch granules and protein bodies for bulk nutrient storage.^[23] Genetic regulation of endosperm composition involves tissue-specific genes and epigenetic mechanisms like genomic imprinting. Endosperm-specific genes, such as those encoding soluble starch synthases (e.g., SSI, SSIIa, SSIIIa in cereals like rice and maize), drive amylopectin synthesis by extending glucan chains in multi-enzyme complexes, with isoforms like OsSSIIa (AF419099) preferentially expressed during grain filling.^[22] Genomic imprinting establishes parent-of-origin effects, where paternally expressed genes (e.g., via reduced maternal methylation) promote endosperm proliferation and nutrient uptake to favor paternal fitness, while maternally expressed genes repress overgrowth to protect maternal resources and prevent excessive seed demand.^[10] This conflict, mediated by DNA methylation and Polycomb proteins, ensures balanced development.^[10] Analytical methods reveal endosperm composition and ploidy through targeted techniques. Histochemical staining, such as iodine-potassium iodide (0.1% I₂ in 1% KI), detects starch by producing blue-black complexes with amylose and amylopectin, distinguishing high-starch wild-type endosperms from mutants with reduced accumulation.^[24] Genomic studies, including RNA-seq, profile expression dynamics; for example, in durum wheat endosperm, RNA-seq across developmental stages (6-30 days post-anthesis) identifies differentially expressed genes (e.g., 29,925 DEGs) in starch biosynthesis pathways like AGPase and GBSS1, yielding datasets for functional annotation.^[25] These approaches, combined with flow cytometry for ploidy confirmation, provide insights into genetic and biochemical variations without relying on embryo nutrition details.

Evolutionary Origins

In Angiosperms

The endosperm in angiosperms originated approximately 140–130 million years ago during the Early Cretaceous, coinciding with the emergence of double fertilization as a key innovation that enabled efficient resource allocation to support embryo development, though molecular clock estimates suggest potentially earlier origins in the late Triassic.^[26]^[27] This process involves the fusion of one sperm cell with the egg to form the diploid embryo and another with the central cell to produce the endosperm, marking a departure from single fertilization in earlier seed plants. Phylogenetically, endosperm is universally present across all angiosperms, from basal lineages to derived clades, but is absent in gymnosperms, underscoring its role as a defining synapomorphy of flowering plants. Fossil evidence from early angiosperm ovules and pollen, such as those from Archaefructus dated to about 125 million years ago, supports the inference of endosperm development in these primitive forms, though direct preservation of internal tissues is rare. The typically triploid nature of endosperm in most angiosperms provides adaptive advantages by resolving parental conflict over resource provisioning, where the 2:1 maternal-to-paternal genome ratio balances maternal control of seed investment with paternal promotion of offspring fitness and growth.^[28] This genomic imbalance mitigates excessive paternal demands that could deplete maternal resources across multiple seeds, enhancing overall reproductive success. Note that diploid endosperm occurs in some basal angiosperms, altering the ratio to 1:1. Key evolutionary hypotheses trace the origin of the second fertilization event to an ancestral condition involving a supernumerary embryo, as proposed by Friedman, who suggested that this second embryo evolved into the specialized nutritive endosperm tissue.^[29] Complementing this, Pien and Grossniklaus explored the evolution of genomic imprinting in endosperm, positing that parent-of-origin gene silencing arose to regulate dosage and resolve conflicts in the (typically) triploid genome, facilitating stable seed development.^[30] Within angiosperm clades, endosperm development transitioned from the ancestral cellular type—characterized by early cell wall formation after each nuclear division—to the derived nuclear (free-nuclear) type, where multiple divisions occur without walls, particularly evident in eudicots following diversification in the late Cretaceous.^[18] This shift, occurring independently in monocots and eudicots, reflects adaptations to varying seed sizes and nutritional strategies during the post-Cretaceous radiation of flowering plants.^[18]

Comparisons with Other Seed Plants

Gymnosperms lack a true endosperm, relying instead on a haploid, multicellular female gametophyte for embryo nourishment. In species such as Pinus (pines), the female gametophyte develops storage cells filled with nutrients prior to fertilization and directly supplies the growing embryo after zygote formation, without involving a second fertilization event.^[31] This contrasts with angiosperms, where the endosperm forms post-fertilization as a dedicated (typically) triploid tissue resulting from double fertilization, providing a more immediate and genetically balanced nutrient source for the embryo.^[2] The endosperm in angiosperms is evolutionarily homologous to the female gametophyte of gymnosperms, representing a modified structure that incorporates paternal genetic contribution via double fertilization to enhance post-zygotic provisioning. This innovation likely arose around 140 million years ago during the Early Cretaceous, replacing the pre-formed haploid storage tissue of gymnosperms and enabling faster embryo development in a more enclosed seed environment.^[2] In non-seed plants, such as ferns, no endosperm exists; the haploid prothallus (gametophyte) serves as the nutritive structure, providing water and nutrients to the developing embryo until the sporophyte becomes independent.^[32] Among basal angiosperms, like Amborella trichopoda, endosperm development retains primitive cellular patterns, such as bipolar division, while still forming the typical triploid tissue from a seven-celled female gametophyte.^[33] In some angiosperms, endosperm is reduced or absent, with perisperm—a diploid maternal tissue derived from the nucellus—serving as a functional equivalent for nutrient storage. For example, in sugar beet (Beta vulgaris), the persistent nucellus develops into perisperm that supports the embryo when endosperm is minimal.^[34] Fossil records from Mesozoic deposits document the transition from gymnosperm-like ovules, featuring exposed female gametophytes, to enclosed angiosperm ovules with internalized endosperm formation, highlighting this as a pivotal innovation contributing to angiosperm diversification and dominance during the Cretaceous.^[35]

Functions in Seed Biology

Nutrition and Support for Embryo

The endosperm serves as the primary nutritional tissue in many angiosperm seeds, particularly those with persistent endosperm, supplying essential metabolites to the developing embryo through coordinated uptake, synthesis, and transfer processes.^[36] During early seed development, nutrients such as sucrose are mobilized from the maternal phloem into the endosperm via specialized transporters, where they are converted into storage reserves like starch and proteins before being exported to the embryo as hexoses.^[37] This transfer occurs primarily through the basal endosperm transfer layer (BETL), where hexose transporters like ZmSWEET4c facilitate the release of sugars to sustain embryo growth.^[38] In addition to nutrient provision, the endosperm provides physical support by acting as a scaffold and barrier within the seed, filling available space to protect the embryo from mechanical stress and maintain turgor pressure.^[39] This structural role is particularly evident in seeds of fleshy fruits, where the expanding endosperm helps regulate internal pressure and prevents embryo displacement during fruit maturation.^[40] The endosperm's nutritional activity begins during the syncytial stage of embryogenesis, when free nuclei proliferate rapidly to establish a nutrient sink, and continues as cellularization occurs, allowing persistent storage accumulation in mature seeds.^[41] Nutrient transfer to the embryo peaks during mid-embryogenesis, with reserves remaining available until degradation initiates at the onset of germination.^[11] Regulatory mechanisms involve endosperm-specific gene expression, such as that of ADP-glucose pyrophosphorylase (AGPase), which catalyzes the synthesis of ADP-glucose for starch production and is upregulated by signals like abscisic acid (ABA) to promote reserve accumulation.^[22] Hormonal cues, including ABA, coordinate these processes by enhancing transporter activity and metabolic pathways in response to developmental cues.^[22] In maize (Zea mays), the endosperm constitutes approximately 85% of the kernel's dry weight and supplies over 80% of the starch that ultimately supports embryo development, highlighting its critical role; disruptions in endosperm function, such as impaired nutrient mobilization, often result in seed abortion.^[42]^[43] In contrast, in non-endospermic seeds such as beans and peas, the endosperm is largely absorbed by the cotyledons during development, transferring its nutritional role to the cotyledons for embryo support and germination.^[3]

Role in Germination and Seed Viability

In seeds with persistent endosperm, the endosperm serves as a critical regulator of dormancy during seed germination, acting as a physical and biochemical barrier that prevents premature radicle emergence until environmental conditions are favorable. In Arabidopsis thaliana, endosperm rupture is a necessary step for radicle protrusion, where the endosperm imposes mechanical restraint on the embryo, requiring weakening through enzymatic degradation to allow growth. This process is modulated by abscisic acid (ABA) signaling within the endosperm, which inhibits cell wall loosening and maintains dormancy by promoting ABI5 accumulation in the embryo, thereby blocking the transition to post-germination growth. In cereals, the endosperm similarly enforces dormancy via ABA-mediated suppression of hydrolytic enzyme production in the aleurone layer, counteracting gibberellin (GA) signals from the embryo to delay reserve mobilization. Seed viability is closely tied to the endosperm's capacity for desiccation tolerance, particularly in orthodox seeds that can withstand drying to low moisture levels (around 5-10%) without losing germination potential. Late embryogenesis abundant (LEA) proteins accumulate in the endosperm during maturation, stabilizing cellular structures and preventing protein denaturation under desiccation stress, which enhances long-term storage viability. In contrast, recalcitrant seeds, which lack robust endosperm desiccation tolerance, exhibit reduced LEA protein expression and fail to maintain viability below 20-30% moisture, leading to rapid deterioration. The quality of endosperm reserves, such as starch and proteins, further influences longevity, with higher accumulation correlating to extended viability periods in orthodox species like wheat and maize. Upon imbibition, the endosperm mobilizes nutrients to support early seedling growth, primarily through the aleurone layer in cereals, which responds to GA diffusion from the embryo by secreting hydrolytic enzymes. GA induces expression of alpha-amylase in aleurone cells, hydrolyzing endosperm starch into simple sugars that are transported to the embryo, with enzyme activity increasing significantly within 24-48 hours post-imbibition and peaking around 3-7 days later in barley.^[44] This coordinated breakdown ensures efficient resource allocation, while proteases and other enzymes degrade storage proteins, providing amino acids for de novo synthesis during germination. Genetic controls in the endosperm contribute to dormancy timing through imprinting mechanisms, where maternally expressed alleles of genes like DELAY OF GERMINATION 1 (DOG1) in the triploid endosperm impose delays in germination. DOG1 imprinting leads to preferential maternal expression, enhancing ABA sensitivity and endosperm weakening resistance, thereby regulating dormancy depth across generations in Arabidopsis. This endosperm-specific regulation integrates with embryonic signals to fine-tune germination responses. Environmental factors influence endosperm function during germination, with water uptake causing swelling that initiates metabolic reactivation and enzyme secretion. Drought or high temperatures can impair viability by disrupting endosperm desiccation tolerance, reducing LEA protein efficacy and accelerating reserve degradation, while optimal temperatures (15-25°C) promote GA-ABA balance for timely endosperm rupture.

Economic and Applied Aspects

In Cereal Grains

In cereal grains, the endosperm is predominantly cellular, featuring a peripheral aleurone layer of living cells that surround a central starchy endosperm core. The aleurone layer remains viable during seed maturation, while starchy endosperm cells undergo programmed cell death to facilitate nutrient storage. The starchy core typically contains 70-80% starch, primarily as amylose and amylopectin, alongside protein bodies that store storage proteins. In maize, these protein bodies accumulate zeins, the major storage proteins, which form within the endoplasmic reticulum and contribute to the endosperm's opacity and texture.^[23]^[45]^[46]^[47] Major cereal crops rely on the endosperm as their primary edible component. In wheat, the endosperm constitutes approximately 83% of the kernel weight and serves as the source for white flour production through milling. Rice processing involves polishing to remove the bran and germ layers, exposing the starchy endosperm for consumption as white rice. Maize endosperm, rich in zeins, has been improved via the opaque-2 mutant, which reduces zein synthesis and increases lysine and tryptophan levels, enhancing overall protein quality in quality protein maize varieties.^[48]^[49]^[50] Milling processes separate the endosperm from the pericarp (bran) and embryo, yielding refined products dominated by the endosperm's composition. This results in a nutritional profile high in carbohydrates (primarily starch) but low in fiber, vitamins, and minerals compared to whole grains, as the bran and germ contain most of these nutrients. Whole grain retention mitigates fiber loss, supporting digestive health and reducing chronic disease risk.^[51]^[52]^[53] Breeding efforts have targeted endosperm traits to boost yield and quality, exemplified by the Green Revolution's semi-dwarf varieties in wheat and rice, which allocate more resources to grain filling under high nitrogen inputs. These dwarfing genes, such as Rht in wheat and sd1 in rice, increased endosperm biomass and global production. However, wheat endosperm's high gluten content—comprising about 85% of its proteins as gliadins and glutenins—poses challenges for individuals with celiac disease, triggering autoimmune responses.^[54]^[55]^[56] Cereals provide about 43-45% of global human caloric intake, with the endosperm forming the bulk of this edible portion according to recent FAO assessments. This underscores the endosperm's role as a staple food source, supporting food security amid population growth.^[57]^[58]

Industrial and Biotechnological Uses

Starch extracted from corn endosperm serves as a primary feedstock for industrial applications, particularly in biofuel production where it is fermented into ethanol. In the United States, ethanol production from corn starch reached approximately 16.2 billion gallons in 2024, supporting renewable energy goals and reducing reliance on fossil fuels.^[59] Additionally, corn endosperm-derived starch is widely used in adhesives for its binding properties in paper products and glues, as well as in textiles as a sizing agent to enhance yarn strength and reduce friction during weaving.^[60] From wheat endosperm, gluten is isolated and applied as a vital additive in baking aids to fortify low-protein flours, improving dough elasticity and volume in industrial bread and pasta production.^[61] In biotechnological contexts, cereal endosperm functions as a natural bioreactor for recombinant protein production, leveraging its storage capacity and isolation from microbial contaminants. For instance, transient expression systems in barley endosperm enable rapid, high-yield production of therapeutic proteins like antimicrobial peptides, with stable inheritance demonstrated across multiple generations in field trials.^[62] CRISPR/Cas9 editing of prolamin genes in rice endosperm has been used to reduce 13 kDa prolamin levels and alter seed protein composition, with studies since 2020 demonstrating viable knockout lines.^[63] Endosperm-derived polysaccharides find pharmaceutical applications in controlled drug delivery systems. Starch-based hydrogels from corn endosperm exhibit tunable swelling and degradation properties, allowing targeted release of therapeutics in the colon for treatments like inflammatory bowel disease.^[64] Beta-glucans extracted from oat endosperm serve as nutraceuticals with immunomodulatory effects, approved for cholesterol reduction and blood glucose management, and are incorporated into formulations for cancer adjunct therapy and wound healing due to their ability to activate immune responses.^[65] Sustainability challenges in endosperm utilization include the environmental footprint of corn monoculture, which consumes vast water resources—approximately 3,000 gallons per bushel (with estimates ranging up to 10,000 gallons in high-evaporation regions)—contributing to groundwater depletion and ecosystem disruption in the U.S. Corn Belt.^[66] As an alternative, endosperm-derived bioplastics from corn starch offer a petroleum-independent option, biodegrading faster than conventional plastics and reducing carbon emissions by 60-80% in packaging applications, though scalability depends on sustainable sourcing to avoid exacerbating monoculture issues.^[67] Recent advancements from 2023 to 2025 highlight endosperm's role in synthetic biology for vaccine production. Rice endosperm has been engineered to express SARS-CoV-2 spike protein S1 subunits, yielding up to 0.282 mg per gram (282 μg/g) of dry seed weight in stable transgenic lines, enabling low-cost, scalable COVID-19 subunit vaccines suitable for developing countries with demonstrated immunogenicity in animal models.^[68]

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In most flowering plants, the endosperm is triploid because of a doubled maternal genetic contribution in the central cell.
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Endosperm - an overview | ScienceDirect Topics
The endosperm is a primary constituent of mature seeds in rice as well as in other cereal crops, serving as the major storage reserve of starch.Missing: minerals | Show results with:minerals
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Starch biosynthesis in cereal endosperms: An updated review over ...
Sep 2, 2021 · Amylose in cereal endosperms is synthesized by granule-bound starch synthase I (GBSSI), which is encoded by the Waxy (Wx) gene (Shure et al., ...
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The aleurone layer of cereal grains: Development, genetic ...
Feb 12, 2025 · Aleurone cells maintain viability throughout seed development, whereas starchy endosperm cells undergo programmed cell death during maturation.
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Sugary Endosperm is Modulated by Starch Branching Enzyme IIa in ...
Jul 20, 2017 · Each seed was stained with iodine solution [0.1% (w/v) I2 and 1% (w/v) KI] to detect starch. Scanning Electron Microscopy. Seeds and starch ...Missing: histochemical | Show results with:histochemical
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Gene expression profile of the developing endosperm in durum ...
Jul 18, 2023 · Mutation of the plastidial alpha-glucan phosphorylase gene in rice affects the synthesis and structure of starch in the endosperm. Plant ...
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Major evolutionary trends in the angiosperm fossil record - PNAS
By Paleocene and Eocene time, there are several evidences in the fossil record of bilateral flowers. This evolution probably began during the Upper Cretaceous.
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Endosperm triploidy has a selective advantage during ongoing ...
It has been argued that a diploid 1m : 1p endosperm was ancestral and the 2m : 1p endosperm evolved after parental conflict, to improve maternal control over ...
[25]
Developmental and evolutionary hypotheses for the origin of double ...
Working with Lilium and Fritillaria, Nawaschin and Guignard were able to document the participation of the second sperm of a pollen tube in a fusion event with ...
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Genomic Imprinting During Seed Development - ScienceDirect.com
Genomic imprinting allows parent-of-origin specific control over gene expression. Although imprinted genes (or entire chromosomes) are homologous sequences ...
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Pinus: pine trees – Inanimate Life - Milne Publishing
Pines reproduce by seed, a multigenerational unit, which in the case of conifers contains both an embryo and the female gametophyte that produced the egg that ...
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Fern Reproduction - Penn Arts & Sciences
The embryonic plant depends upon the prothallus for water and nutrients. As the embryo grows and develops into a mature diploid plant the prothallus dies.
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Developmental evolution of endosperm in basal angiosperms
We provide detailed descriptions of endosperm development in Amborella, Nuphar (Nymphaeales), and Illicium (Illiciales) and compare patterns within an explicit ...
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Hybridity has a greater effect than paternal genome dosage on ...
Jun 15, 2018 · In sugar beet seed development, the maternal nucellus is not fully digested during maturation and leads to the perisperm tissue which ...
[31]
Fossil evidence and phylogeny: the age of major angiosperm clades ...
Oct 1, 2004 · The fossil record has played an important role in the history of evolutionary thought, has aided the determination of key relationships ...
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Embryo-Endosperm Interaction and Its Agronomic Relevance to ...
Nov 30, 2020 · The endosperm consists of aleurone (white) and starchy endosperm (pink) that are packed with starch granules and protein bodies, respectively.
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A Cascade of Sequentially Expressed Sucrose Transporters in the ...
This cascade of sequentially expressed SWEETs provides the feeding pathway for the plant embryo, an important feature for yield potential.
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Plant SWEETs: from sugar transport to plant–pathogen interaction ...
This hexose transporter is expressed in the basal endosperm transfer layer region and is responsible for transferring hexoses to sustain development of ...
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The Metabolic Role of the Legume Endosperm: A Noninvasive ...
The endosperm provides a temporary source of nutrition, permits space for embryo growth, and acts as a buffer between the maternal organism and its offspring.
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the impact of the endosperm and other extra-embryonic seed tissues ...
Jan 14, 2020 · Nutrients are then re-exported from the endosperm for absorption by the embryo, a process facilitated by the ephemeral basal region of embryo ...
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Transcriptome Analysis of Proliferating Arabidopsis Endosperm ...
During the early stages of seed development, Arabidopsis (Arabidopsis thaliana) endosperm is syncytial and proliferates rapidly through repeated rounds of ...
[38]
Phenotypic and Transcriptional Analysis of Divergently Selected ...
Endosperm is a major storage organ in maize (Zea mays), accounting for roughly 85% of the dry seed weight and containing over 80% of total seed starch and about ...
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Postzygotic reproductive isolation established in the endosperm
The endosperm is a developmental innovation of angiosperms that supports embryo growth and germination. Aside from this essential reproductive function, the ...
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3D Electron Microscopy Gives a Clue: Maize Zein Bodies Bud From ...
Jun 11, 2020 · Zeins are the main storage proteins in maize seed endosperm, and the onset of zein synthesis in young seeds challenges the endomembrane ...
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Genetic control of cereal kernel texture: Towards a maize model
Sep 12, 2025 · In cereal endosperm, starch is composed of amylose and amylopectin. Among diverse parameters describing starch, amylose content (AC) is the most ...
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Cereal seed storage proteins: structures, properties and role in grain ...
In maize, the protein bodies in the sub‐aleurone and outer parts of the endosperm are enriched in γ‐ and β‐zeins and low in α‐zeins, the latter being more ...Missing: core | Show results with:core
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A Kernel of Wheat | Learn
Sep 29, 2013 · Endosperm. About 83 percent of the kernel weight and the source of white flour. Bran. About 14 ½ percent of the kernel weight. Bran is ...
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Rice Polishing Guide for Sake - BrewSake.org
The purpose of milling is to remove the husk (hull) of the grain, followed by up to 10% of the outer layer (bran, germ) from the brown rice grains.
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Expression profile of protein fractions in the developing kernel of ...
Jan 28, 2021 · Endosperm modification of opaque-2 had led to the development of Quality Protein Maize (QPM), which has higher protein quality along with ...
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Cereal by-products as an important functional ingredient: effect of ...
Oct 24, 2018 · Cereal grains are usually milled to remove the fibrous bran, during milling, bran which is separated from the starchy endosperm of the grain is ...
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Improving cereal grain carbohydrates for diet and health - PMC - NIH
Starch and cell wall polysaccharides (dietary fibre) of cereal grains contribute to the health benefits associated with the consumption of whole grain cereal ...
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(PDF) Nutritional aspects of cereals - ResearchGate
Aug 6, 2025 · Cereals are low in sodium and are a good source of potassium, in common with most plant foods. Wholegrain cereals also contain considerable amounts of iron, ...
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Generating high-yielding varieties by genetic manipulation of plant ...
In this review, we highlight findings on the dwarfing genes used in the green revolution and recent developments in the molecular breeding of semi-dwarf ...
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Reinventing a sustainable Green Revolution by breeding and ...
The agricultural Green Revolution gave rise to semi-dwarf varieties that increased wheat and rice yields under nitrogen-intensive and high-density planting ...
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Next-generation therapies for celiac disease: The gluten-targeted ...
Gluten, which corresponds to about 85% of the total proteins in the wheat endosperm, is comprised of two types of proteins, gliadins and glutenins (Wieser, 2007) ...
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https://pmc.ncbi.nlm.nih.gov/articles/PMC4064937/
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[PDF] 3 Cereals - FAO Knowledge Repository
Consequently, and because cereals represent a significant portion of global consumption (about 45% of calories), their contribution to overall calorie loss and ...
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[PDF] Economic Impact of the U.S. Ethanol Industry
Feb 17, 2025 · Ethanol production approached a record 16.1 billion gallons (bg) in 2024, in response to solid domestic consumption and exports that ...Missing: endosperm starch
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Corn Starch: Quality and Quantity Improvement for Industrial Uses
Dec 28, 2021 · Therefore, it is mainly used by food companies as a stabilizer and thickener. Moreover, it is used in the textile, adhesive, corrugating, and ...
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Protein Characteristics that Affect the Quality of Vital Wheat Gluten to ...
Mar 2, 2017 · In bakery products, vital wheat gluten is used to fortify flour with lower protein content and of less than desirable quality or is inadequate ...
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Long-Lasting Stable Expression of Human LL-37 Antimicrobial ... - NIH
The use of barley grains as a desirable bioreactor has been developed for the production of human therapeutic proteins because the endosperm of barley grain is ...
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Multiplex CRISPR/Cas9 Editing of Rice Prolamin and GluA Glutelin ...
Jul 31, 2025 · To enhance seed quality, CRISPR/Cas9 genome editing was applied to modify SSP composition by targeting genes encoding 13 kDa prolamins and type ...Missing: trials | Show results with:trials
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Starch hydrogels as targeted colonic drug delivery vehicles
Aug 1, 2022 · In this work we show how starch hydrogel bulk structural and molecular level parameters influence their properties as drug delivery platforms.
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Pharmaceutical application of beta-glucan: a comprehensive overview
Jun 20, 2025 · Their immunomodulatory properties make them valuable in various fields, including cancer treatment, vaccine development, wound healing, and ...Role Of Beta-Glucan · Beta-Glucan In Drug Delivery... · Cancer Treatment And...
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How Does Corn Farming Work and Is It Bad for the Environment?
Mar 1, 2022 · Industrialized corn farms use a lot of water, which is draining groundwater in the middle of the U.S. The excessive fertilizer from corn farms ...Missing: endosperm | Show results with:endosperm
[62]
Development and characterization of starch bioplastics as a ... - Nature
May 1, 2025 · This study seeks to bridge that gap by formulating and characterizing biodegradable plastics using corn starch, potato starch, and their blends, ...
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Rice‐derived SARS‐CoV‐2 glycoprotein S1 subunit vaccine elicits ...
Apr 4, 2025 · These findings suggest that rS1 protein offers a promising approach for affordable COVID-19 subunit vaccine production, and this strategy can be ...