double fertilization process

Nat. They develop within the microsporangia, or pollen sacs, of the anthers on the stamens. In angiosperms, one sperm fertilizes the egg to form the 2n zygote, and the other sperm fertilizes the central cell to form the 3n endosperm. The ovary, surrounding the ovules, develops into the fruit, which protects the seeds and may function to disperse them.[1]. heterostyly: the condition of having unequal male (anther) and female (stigma) reproductive organs cross-pollination: fertilization by the transfer of pollen from an anther of one plant to a stigma of another self-pollination: pollination of a flower by its own pollen in a flower that has both stamens and a pistil Pollination: An Introduction It is controversial whether this requires active transport or is solely based on cytoplasmic flow associated with burst of both pollen tube and receptive synergid cell and/or the architecture of the egg apparatus. A sex recognition glycoprotein is encoded by the plus mating-type gene fus1 of Chlamydomonas reinhardtii. Rev. Abstract During the double fertilization process, pollen tubes deliver two sperm cells to an ovule containing the female gametes. (2011). (2007). Development 136, 32793288. Identify your study strength and weaknesses. If you're behind a web filter, please make sure that the domains *.kastatic.org and *.kasandbox.org are unblocked. The terminal cell also divides, producing a globular-shaped pro-embryo that develops into an embryo forming the baby plant. 101, 712. The immobile sperm cells are delivered by the pollen tube toward the ovule harboring the . Double fertilization occurs only in angiosperms (flowering plants) inside the female structures in the flower. The microspores, or the pollen, contain two cells: the pollen tube cell and the generative cell. The essential role of calcium ion in pollen germination and pollen tube growth. (2003). During this process, if the generative cell has not already split into two cells, it now divides to form two sperm cells. Remember that the gametophyte of gymnosperms is also a pollen grain that produces two sperm cells. doi: 10.1371/journal.pgen.1003933, Liu, J., Zhong, S., Guo, X., Hao, L., Wei, X., Huang, Q., et al. In vitro fertilization (IVF) is a treatment for infertility or genetic problems. The other sperm fuses with the two polar nuclei, resulting in a triploid cell. J. Upon landing on the female cone, the tube cell of the pollen forms the pollen tube, through which the generative cell migrates towards the female gametophyte through the micropyle. doi: 10.1371/journal.pgen.1000430, Calder, G. M., Franklin-Tong, V. E., Shaw, P. J., and Drbak, B. K. (1997). 270 lessons doi: 10.1104/pp.113.222950, Stratford, S., Barne, W., Hohorst, D. L., Sagert, J. G., Cotter, R., Golubiewski, A., et al. The product of the second fertilization, the endosperm, represents a highly nutritious source for the developing embryo that increases its survival probabilities. Hence pollen tubes maturate during their growth through the sporophytic tissue and thereby become competent for fertilization (Leydon et al., 2013, 2014). Create your account. After fertilization, the zygote divides to form an upper terminal cell and a lower basal cell. Nature 459, 992995. Depending on the plant species, the endosperm might be transient. Nat. doi: 10.1104/pp.113.231274, Hamamura, Y., Saito, C., Awai, C., Kurihara, D., Miyawaki, A., Nakagawa, T., et al. doi: 10.1104/pp.107.098236, Schopfer, C. R. (1999). Genetic analysis of female gametophyte development and function. Identification and characterization of components of a putative petunia S-locus F-box-containing E3 ligase complex involved in S-RNase-based self-incompatibility. (2004). doi: 10.1038/ng1694, Ogawa, M., Kay, P., Wilson, S., and Swain, S. M. (2009). In the typical flowering plant, seeds are formed from bodies calledovules, which arecontained in theovary, or basal part of the female plant structure, thepistil of a flower. 14, 588593. - Definition & Food Examples, Working Scholars Bringing Tuition-Free College to the Community. Out of the two sperm cells, one fertilizes the egg cell to produce a zygote, and the remaining one fuses with two polar nuclei producing endosperm. Pollen grains are transported to the female part of another flower through pollination. A., Haring, V., Ebert, P. R., Anderson, M. A., Simpson, R. J., Sakiyama, F., et al. doi: 10.1093/aob/mcs143, Nodine, M. D., and Bartel, D. P. (2012). The major function of the filiform apparatus may thus be to considerably increase the micropylar surface of the synergid cells, which represent glandular cells of the egg apparatus. 108, 727737. Thus, the endosperm is the direct nutrient source for monocotyledons embryos. 110, 383404. Initially, papilla cells export Ca2+cyto by the auto-inhibited Ca2+-ATPase13 (ACA13) at the pollen grain adhesion site (Iwano et al., 2004, 2014). A positive signal from the fertilization of the egg cell sets off endosperm proliferation in angiosperm embryogenesis. Res. It develops from the primary endosperm cell during double fertilization. 9:e1003933. The haploid sperm and haploid egg combine to form a diploid zygote, the process being called syngamy, while the other sperm and the two haploid polar nuclei of the large central cell of the megagametophyte form a triploid nucleus (triple fusion). The central cell plays a critical role in pollen tube guidance in Arabidopsis. Earn points, unlock badges and level up while studying. We'll describe the process of double fertilization and its advantages, unique to flowering plants, that originate this food-storage tissue. Plant Mol. Sperm entry is sufficient to trigger division of the central cell but the paternal genome is required for endosperm development in Arabidopsis. This is called a double fertilization. These cookies do not store any personal information. Directional guidance of Nicotiana alata pollen tubes in vitro and on the stigma. The lower end of the embryo sac consists of the haploid egg cell positioned in the middle of two other haploid cells, called synergids. 48, 5472. Is it the same thing as triple fusion? Molecular genetics, physiology and biology of self-incompatibility in Brassicaceae. (2012) and Dresselhaus and Franklin-Tong (2013). These changes in Ca2+cyto level are essential for sperm delivery and are depending on FER and LRE activity, respectively. Nature 434, 234438. Bot. Biol. J. Integr. You now know that gymnosperms seeds are naked (exposed on the surface of cones bracts or scales) while angiosperms have seeds enclosed by a fruit. Try refreshing the page, or contact customer support. A calcium dialog mediated by the FERONIA signal transduction pathway controls plant sperm delivery. Double fertilization in Arabidopsis thaliana involves a polyspermy block on the egg but not the central cell. Mol. Plant Cell 21, 26552671. Curr. Double fertilization is a process of fertilization characterized by the fusion of a female gametophyte with two male gametes. The microspores, or the pollen, contain two cells: the pollen tube cell and the generative cell. Some reports suggest that fertilization of the egg cell is preferred, which was demonstrated, for example, in mutants of CYCLIN DEPENDENT KINASE A1 (CDKA;1), which generate only one sperm-like germ cell (Iwakawa et al., 2006; Nowack et al., 2006). doi: 10.1016/0092-8674(95)90427-1, Zhou, L., Fu, Y., and Yang, Z. doi: 10.1242/dev.00458, Ingouff, M., Sakata, T., Li, J., Sprunck, S., Dresselhaus, T., and Berger, F. (2009). Only one of the four resulting megaspores survives. 8:e1000388. The pollen tube cell grows into a pollen tube through which the generative cell travels. Plant Physiol. of the users don't pass the Double Fertilization quiz! At full maturity the nucellus cell (NC) layer surrounding the developing embryo sac is flattened between inner integument (II) and female gametophyte cells. Explore the definition of reproductive structures in angiosperms, pollination, and the process of double fertilization. Flowers are the reproductive structures of angiosperms. Plant Cell 22, 17021715. The immunoglobulin superfamily protein Izumo is required for sperm to fuse with eggs. (2014). (D) DIC microscopic image of a mature female gametophyte surrounded by the maternal sporophytic tissues of the ovule. RALF, a 5-kDa ubiquitous polypeptide in plants, arrests root growth and development. Science 287, 321324. (2000). Biol. Arabidopsis tetraspanins were shown to form homo- and heterodimers, but so far functional studies are missing. Initial attempts enabling ovules to attract pollen tubes from unrelated plant families have been successful (Mrton et al., 2012). After pollination brings a pollen grain to a flower's female reproductive structure (carpel), a pollen tube releases two sperm cells. Central cell-derived peptides regulate early embryo patterning in flowering plants. Types of Blood Cells With Their Structure, and Functions, The Main Parts of a Plant With Their Functions, Parts of a Flower With Their Structure and Functions, Parts of a Leaf With Their Structure and Functions, Plant Cell: Parts and Structure With Functions, On reaching the stigma, the tube cell forms a pollen tube through the. In fer mutant, for example, pollen tube growth continues around the synergid cells, which must be associated with mechanical stress but does not induce synergid cell death (Escobar-Restrepo et al., 2007). Bot. This occurs inside the ovule that is enclosed by the ovary (in the flower). Science 317, 656660. Science 293, 14801483. 1, 96113. (c) In the third stage, the growing embryo runs out of room and starts to bend. The synergid cells represent the main source for chemo-attractants required for micropylar pollen tube guidance. In gymnosperms, pollination involves pollen transfer from the male cone to the female cone. Double fertilization happens when pollen is deposited from the anther (male) to the flowers stigma (female). Acta 1833, 15731581. Both genes are expressed in synergid cells and show a similar pollen tube overgrowth phenotype (Capron et al., 2008; Kessler et al., 2010). A compatible pollen tube expresses the S-locus F-box protein (SLF), which leads to the degradation of the S-RNase (Hua and Kao, 2006; Kubo et al., 2010), while in incompatible interactions intact S-RNase degrades RNAs resulting, for example, in the disruption of the actin cytoskeleton and other cellular processes (Liu et al., 2007; Roldn et al., 2012). doi: 10.1073/pnas.2533800100, Koltunow, A. M., and Grossniklaus, U. [9] The more primitive process of double fertilization in gymnosperms results in two diploid nuclei enclosed in the same egg cell. During pollen tube growth the tip needs to modulate the surrounding cell wall of stylar cells enabling its penetration through the extra-cellular space, most likely by interaction with extensin-like proteins and arabinogalactan proteins as well as the secretion of cell wall softening enzymes and inhibitors such as polygalacturonases and pectin methylesterase inhibitors (Cosgrove et al., 1997; Grobe et al., 1999; Stratford et al., 2001; Ogawa et al., 2009; Nguema-Ona et al., 2012; Woriedh et al., 2013). Evidence suggests that the synergids secrete chemical signals to guide the pollen tube near the egg cell. doi: 10.1146/annurev-genet-110711-155527. 22, 11941198. 131, 814823. Other seeds, such as those ofbeets, contain both perisperm and endosperm. If you're seeing this message, it means we're having trouble loading external resources on our website. Ethylene signaling is required for synergid degeneration and the establishment of a pollen tube block. doi: 10.1016/j.tcb.2009.12.007, Woriedh, M., Wolf, S., Mrton, M. L., Hinze, A., Gahrtz, M., Becker, D., et al. An electron microscopic study of the mature megagametophyte in Zea mays. The male gametophytes, or microgametophytes, that participate in double fertilization are contained within pollen grains. From the moment of germination until sperm discharge the pollen grain/tube communicates with at least five different sporophytic and three different gametophytic cell types to successfully accomplish fertilization (Palanivelu and Tsukamoto, 2012). The ovule is connected to the septum (SE, yellow) containing the transmitting tract (TT, blue) by the funiculus (F, petrol) and surrounded by the carpel tissue (C) (green). This page titled 40.5D: Double Fertilization in Plants is shared under a not declared license and was authored, remixed, and/or curated by Boundless. After arrival at the surface of the ovule, the pollen tube reaches the last phase of its journey, which is known as micropylar pollen tube guidance. doi: 10.1091/mbc.E02-12-0790, Miyado, K., Yamada, G., Yamada, S., Hasuwa, H., Nakamura, Y., Ryu, F., et al. Two closely related homologs of FER, ANXUR1 (ANX1), and ANX2 were reported to be involved in the timing of pollen tube burst or more precisely in the inhibition of pollen tube burst. Near to the micropyle (MY), the two nuclei of the two synergid cells (SY) are shown in red and green. The basal cell also divides, giving rise to the suspensor. doi: 10.1104/pp.014597, Haruta, M., Monshausen, G., Gilroy, S., and Sussman, M. R. (2008). The cells of an embryo sac of an unfertilized ovule are 8 in number and arranged in the form of 3+2+3 (from top to bottom) i.e. Curr. After pollen tube burst inside the female gametophyte, the two sperm cells fuse with the egg and central cell initiating seed development. Due to their rapid molecular evolution it was difficult to identify orthologs in other plant species, but finally the DEFL subgroup CRP810/AtLURE1 of Arabidopsis was discovered to be involved in micropylar pollen tube guidance (Takeuchi and Higashiyama, 2012). The triploid cell grows into the endosperm and will nourish the developing embryo. Misamore, M. J., Gupta, S., and Snell, W. J. Its like a teacher waved a magic wand and did the work for me. doi: 10.1016/j.cub.2012.04.061, Maruyama, D., Hamamura, Y., Takeuchi, H., Susaki, D., Nishimaki, M., Kurihara, D., et al. Ultimately, the embryo and cotyledons fill the seed, at which point, the seed is ready for dispersal. The pollen tube grows on the septum surface toward the funiculus, the tissue connecting the ovule with the septum (Figures 1BD; Supplemental Movies 1, 2). Mol. The terminal cell also divides, giving rise to a globular-shaped proembryo (Figure 2a). doi: 10.1038/33929, Wong, J. L., and Johnson, M. (2010). GEX2 encodes a single-pass transmembrane protein with filamin repeats exposed to the extracellular space. 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It cannot be explained by mechanical breakdown due to an invading pollen tube. doi: 10.1242/dev.040071, Brewbaker, J. L., and Kwack, B. H. (1963). Evolutionary history of the HAP2/GCS1 gene and sexual reproduction in metazoans. The newly formed zygote then undergoes a limited number ofdivisionsand gives rise to an embryo. 8, 6471. Fusion of sperm cell (male gamete) with egg cell (female gamete) to form a diploid zygote. [6], A far more rudimentary form of double fertilization occurs in the sexual reproduction of an order of gymnosperms commonly known as Gnetales. A seed consists of the external seed coat and two internal structures: the embryo and endosperm. 24, 279282. Its synthesizing enzyme gene SR1 is also expressed in the ovule indicating the presence of D-serine (Michard et al., 2011). Later, the pollen tube enters the ovule sac through the micropyle. The central cell surrounds the egg apparatus. In angiosperms, unlike gymnosperms, the endosperm is formed: Which of the following is necessary for a pollen grain to germinate? An error occurred trying to load this video. On the other hand, dicotyledon species (like beans, peanuts, and peas) have embryos with two cotyledons that take over the storage of nutrients. Thus, in gymnosperms, if fertilization does not occur, the resources used for the growth of the female gametophyte might be wasted. doi: 10.1016/j.cub.2012.04.041, Berger, F., Hamamura, Y., Ingouff, M., and Higashiyama, T. (2008). (2013). In flowering plants, fertilization is unique because it involves two pairs of male and female gametes, a process known as double fertilization. Genet. In mammals, CD9-like membrane spanning proteins of the tetraspanin family are located at the plasma membrane of eggs and were shown to be required for gamete fusion (Kaji et al., 2000; Le Naour et al., 2000; Miyado et al., 2000). MAA3 (MAGATAMA3) helicase gene is required for female gametophyte development and pollen tube guidance in Arabidopsis thaliana. (2001). (2011). Annu. Science 330, 796799. Since two fertilization events coincide, it is called double fertilization. Plus, get practice tests, quizzes, and personalized coaching to help you [3], Double fertilization was discovered more than a century ago by Sergei Nawaschin in Kyiv,[4] and Lon Guignard in France. 10:e1001449. The significance of double fertilization in flowering plants is the increased survival probability of the embryo provided by a highly nutritious tissue (endosperm), and to avoid the waste of resources by forming the endosperm only if the egg is fertilized. Create and find flashcards in record time. Female components, which are directly involved in gamete fusion are so far unknown in higher plants. Curr. The male and female . Development 138, 29993009. doi: 10.1007/s00497-013-0222-y, Bayer, M., Nawy, T., Giglione, C., Galli, M., Meinnel, T., and Lukowitz, W. (2009). Pollen hydration and germination appear not to be affected, although only grass pollen tubes are capable of penetrating the style and reach the transmitting tract. Nat. The embryo sac is derived either from one (monosporic) or more (bisporic or tetrasporic) meiotic products (functional megaspores) . They surround the pistil, which represents the female flower organ. In grasses the ovary contains a single ovule and the pollen tube is directly guided toward its surface after leaving the blind ending transmitting tract. GAMETE EXPRESSED 3 (GEX3) is a plasma membrane-localized protein, which is expressed in the unfertilized egg cell. 42, 358363. Supplements. Signalling events regulating seed coat development. The female gametophyte is deeply embedded and thus protected by the maternal sporophytic tissues of the pistil (Figure 1). All angiosperms are characterised by a unique process called double fertilisation. Copyright 2014 Bleckmann, Alter and Dresselhaus. doi: 10.1111/j.1744-7909.2009.00847.x, Keywords: pollen tube, ovule, gamete interaction, cell fusion, signaling, fertilization, polyspermy, Citation: Bleckmann A, Alter S and Dresselhaus T (2014) The beginning of a seed: regulatory mechanisms of double fertilization. What is the advantage of double fertilization? The growth direction of the pollen tube is regulated by the formation of different gradients including water, -amino butyric acid (GABA), calcium and other small molecules such as D-serine. It develops from the primary endosperm cell during double fertilization. - Facts, Addiction & Withdrawal Symptoms, Human Growth Hormones: Uses & Side Effects, Tetramer: Definition, Analysis & Immunology, What Are Complete Proteins? In a transient expression system NTA is directly targeted to the plasma membrane. Science 343, 408411. Paternal control of embryonic patterning in Arabidopsis thaliana. PLoS Biol. However, their presence at the surface of plant gametes and structural homology to mammalian CD9-like proteins suggest that they may possess a similar role during gamete interaction. Pollengrains that land on the receptive upper surface (stigma) of the pistil will germinate, if they are of the same species, and produce pollen tubes, each of which grows down within the style (the upper part of the pistil) toward an ovule. The two sperm cells are connected to each other, likely involving tetraspanins. Biophys. The large cell of the gametophyte will then develop into the endosperm, a nutrient-rich tissue which provides nourishment to the developing embryo. GEX2 localizes to the sperm cell plasma membrane and contains extracellular immunoglobulin-like domains, similar to gamete interaction factors reported in algae and mammals (Misamore et al., 2003; Inoue et al., 2005). Plant Physiol. lip1/2 double mutant pollen reach the funiculus but fail to grow through the micropyle inside the ovule, and the pollen tube shows a reduced attraction toward AtLURE1 (Liu et al., 2013). Out of the two sperm cells, one fertilizes the egg cell and forms a diploid zygote (2n). doi: 10.1242/dev.052928, Barcaccia, G., and Albertini, E. (2013). Embryonic development is suspended after some time; growth resumes only when the seed germinates. However, with the development of the electron microscope, many of the questions were answered. Double fertilization is a key event in the life cycle of angiosperms, but is completely absent in gymnosperms. The microspores, or the pollen, contain two cells: the pollen tube cell and the generative cell. A smaller fraction of flowering plants are wind-pollinated. In diploid (2n) cells, one set of chromosomes comes from the male parent and the other set comes from the female parent. gex2 mutant sperm cells show reduced adhesion to female gametes, likely causing cell fusion failure (Mori et al., 2014). The ovule becomes the __ and the ovary becomes the ___. Psychological Research & Experimental Design, All Teacher Certification Test Prep Courses, Animal Reproduction & Development Overview, Semipermeable Membranes' Role in Cell Communication, Nuclear Envelope: Definition, Function & Structure, Double Fertilization in Angiosperms: Definition & Process, Methionine: Structure, Production & Benefits, Microfilaments: Definition, Function & Structure, Transformed Cells: Definition & Characteristics, The Circulatory, Respiratory, Digestive, Excretory, & Musculoskeletal Systems, High School Chemistry: Homework Help Resource, SAT Subject Test Biology: Practice and Study Guide, Praxis Biology and General Science: Practice and Study Guide, CSET Science Subtest II Chemistry (218): Practice & Study Guide, Introduction to Environmental Science: Help and Review, What Is Albinism? Corrections? Describe the process of double fertilization in plants Double Fertilization After pollen is deposited on the stigma, it must germinate and grow through the style to reach the ovule. 19, 13271331. doi: 10.1126/science.286.5445.1697, Scott, R. J., Armstrong, S. J., Doughty, J., and Spielman, M. (2008).

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