Nucellar embryony (notated Nu+) is a form of seed reproduction that occurs in certain plant species, including many citrus varieties. Nucellar embryony is a type of apomixis, where eventually nucellar embryos from the nucellus tissue of the ovule are formed, independent of meiosis and sexual reproduction. [1] During the development of seeds in plants that possess this genetic trait, the nucellus tissue which surrounds the megagametophyte can produce nucellar cells, also termed initial cells. These additional embryos (polyembryony) are genetically identical to the parent plant, rendering them as clones. By contrast, zygotic seedlings are sexually produced and inherit genetic material from both parents. Most angiosperms reproduce sexually through double fertilization. Different from nucellar embryony, double fertilization occurs via the syngamy of sperm and egg cells, producing a triploid endosperm and a diploid zygotic embryo. In nucellar embryony, embryos are formed asexually from the nucellus tissue. Zygotic and nucellar embryos can occur in the same seed (monoembryony), and a zygotic embryo can divide to produce multiple embryos. [2] The nucellar embryonic initial cells form, divide, and expand. Once the zygotic embryo becomes dominant, the initial cells stop dividing and expanding. Following this stage, the zygotic embryo continues to develop and the initial cells continue to develop as well, forming nucellar embryos. The nucellar embryos generally end up outcompeting the zygotic embryo, rending the zygotic embryo dormant. The polyembryonic seed is then formed by the many adventitious embryos within the ovule [3] (to picture this process, refer to Figure 1). The nucellar embryos produced via apomixis inherit its mother's genetics, making them desirable for citrus propagation, research, and breeding. [4]
Nucellar embryos have also been found in polyembryonic Mango varieties, where generally one of the embryos is zygotic and the rest are nucellar. [5] However, there is little research on Mangos undergoing nucellar embryo development as there has on varieties of citrus.
Nucellar embryony is able to occur within both fertilized and unfertilized ovules. Furthermore, instead of using the endosperm as nutritive tissue, it will utilize the surrounding nucellus tissue for nutrition. [3] For example, the ‘Valencia’ orange undergoes nucellar embryony in both fertilized and unfertilized conditions. [3] But, it has been found that nucellar embryo development, under fertilized or unfertilized conditions, can take place in different positions. [6]
An important component of nucellar embryo development is its changing cell wall thickness. Between nucellar embryo's initial cell stage and its dividing and expanding stage, the cells' wall thickens. [7] This most likely occurs due to callose deposition; callose deposition reduces the permeability of a cell and is usually found in the initial cells about to undergo embryogenesis. [8] The initial cells become enlarged, rounded, and divided. During this stage, the initial cell's cell walls thin out, leaving room for the nucleus to become distinguished.
Many seed plants, including citrus fruits, are self-compatible, meaning that they are able to fertilize themselves. Self-compatibility produces a seedy fruit which may be deemed as undesirable to the citrus industry.
Seedless fruits have been made popular as they are sought-after in the citrus industry. To be seedless, a citrus must exhibit self-incompatibility, another reproductive trait within citrus fruits and many seed plants. Self incompatibility is the phenomena where hermaphroditic plants are not able to produce fertile embryos after self-pollination. [9] Self-incompatibility is regulated by the S-loci; if pollen is rendered incompatible, it is determined by its haploid S genotype, or if its sporophyte is rendered incompatible, it would be determined by its diploid S genotype. This is also termed and associated with parthenocarpy, the production of fruit without fertilization. Self-incompatible fruits are able to undergo parthenocarpy to yield seedless fruits. In citrus specifically, there have been other modes developed to reduce seeding as well: gibberellic acid enhances ovule abortion [10] and copper sulfate has been shown to reduce seed number in fruit. [11] An example is the ‘Afourer’ mandarin that contains a haploid self-incompatibility system and parthenocarpy. Under conditions where cross-pollination is not present, the ‘Afourer’ mandarin produces a seedless fruit by undergoing parthenocarpy. Where cross-pollination is present, gibberellic acid is applied and produces a decreased seeding fruit. [11]
Nucellar embryony is important to the citrus industry, as it allows for the production of uniform rootstock which yields consistent results in fruit production. However, this trait can interfere with progress in cross-breeding; most commercial scion varieties produce mainly nucellar seedlings which do not inherit any of the traits of the "father" plant.
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the full set of genes of their single parent and thus the newly created individual is genetically and physically similar to the parent or an exact clone of the parent. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many eukaryotic organisms including plants, animals, and fungi can also reproduce asexually. In vertebrates, the most common form of asexual reproduction is parthenogenesis, which is typically used as an alternative to sexual reproduction in times when reproductive opportunities are limited. Komodo dragons and some monitor lizards can reproduce asexually.
In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering.
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae, commonly called angiosperms. They include all forbs, grasses and grass-like plants, a vast majority of broad-leaved trees, shrubs and vines, and most aquatic plants. The term "angiosperm" is derived from the Greek words ἀγγεῖον / angeion and σπέρμα / sperma ('seed'), meaning that the seeds are enclosed within a fruit. They are by far the most diverse group of land plants with 64 orders, 416 families, approximately 13,000 known genera and 300,000 known species. Angiosperms were formerly called Magnoliophyta.
In botany, a seed is a plant embryo and food reserve enclosed in a protective outer covering called a seed coat (testa). More generally, the term "seed" means anything that can be sown, which may include seed and husk or tuber. Seeds are the product of the ripened ovule, after the embryo sac is fertilized by sperm from pollen, forming a zygote. The embryo within a seed develops from the zygote and grows within the mother plant to a certain size before growth is halted.
In botany, apomixis is asexual development of seed or embryo without fertilization. However, other definitions include replacement of the seed by a plantlet or replacement of the flower by bulbils.
In botany and horticulture, parthenocarpy is the natural or artificially induced production of fruit without fertilisation of ovules, which makes the fruit seedless. The phenomenon has been observed since ancient times but was first scientifically described by German botanist Fritz Noll in 1902.
In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the integument, forming its outer layer, the nucellus, and the female gametophyte in its center. The female gametophyte — specifically termed a megagametophyte — is also called the embryo sac in angiosperms. The megagametophyte produces an egg cell for the purpose of fertilization. The ovule is a small structure present in the ovary. It is attached to the placenta by a stalk called a funicle. The funicle provides nourishment to the ovule. On the basis of the relative position of micropyle, body of the ovule, chalaza and funicle, there are six types of ovules.
The endosperm is a tissue produced inside the seeds of most of the flowering plants following double fertilization. It is triploid in most species, which may be auxin-driven. It surrounds the embryo and provides nutrition in the form of starch, though it can also contain oils and protein. This can make endosperm a source of nutrition in animal diet. For example, wheat endosperm is ground into flour for bread, while barley endosperm is the main source of sugars for beer production. Other examples of endosperm that forms the bulk of the edible portion are coconut "meat" and coconut "water", and corn. Some plants, such as certain orchids, lack endosperm in their seeds.
Self-incompatibility (SI) is a general name for several genetic mechanisms that prevent self-fertilization in sexually reproducing organisms, and thus encourage outcrossing and allogamy. It is contrasted with separation of sexes among individuals (dioecy), and their various modes of spatial (herkogamy) and temporal (dichogamy) separation.
Plant embryonic development, also plant embryogenesis, is a process that occurs after the fertilization of an ovule to produce a fully developed plant embryo. This is a pertinent stage in the plant life cycle that is followed by dormancy and germination. The zygote produced after fertilization must undergo various cellular divisions and differentiations to become a mature embryo. An end stage embryo has five major components including the shoot apical meristem, hypocotyl, root meristem, root cap, and cotyledons. Unlike the embryonic development in animals, and specifically in humans, plant embryonic development results in an immature form of the plant, lacking most structures like leaves, stems, and reproductive structures. However, both plants and animals including humans, pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.
Gynoecium is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of pistils and is typically surrounded by the pollen-producing reproductive organs, the stamens, collectively called the androecium. The gynoecium is often referred to as the "female" portion of the flower, although rather than directly producing female gametes, the gynoecium produces megaspores, each of which develops into a female gametophyte which then produces egg cells.
Bromelia hieronymi is a species of plant in the family Bromeliaceae native to Argentina, Bolivia and Paraguay. It is one of several plants used by the Wichí people as a fiber for weaving called chaguar. It has anti-inflammatory agents that are secreted by the fruit.
Stenospermocarpy is the biological mechanism that produces parthenocarpy (seedlessness) in some fruits, notably many table grapes.
Double fertilization or double fertilisation is a complex fertilization mechanism of angiosperms. This process involves the fusion of a female gametophyte or megagametophyte, also called the embryonic sac, with two male gametes (sperm). It begins when a pollen grain adheres to the stigmatic surface of the carpel, the female reproductive structure of angiosperm flowers. The pollen grain begins to germinate, forming a pollen tube that penetrates and extends down through the style toward the ovary as it follows chemical signals released by the egg. The tip of the pollen tube then enters the ovary by penetrating through the micropyle opening in the ovule, and releases two sperm into the embryonic sac (megagametophyte).
The Valencia orange is a sweet orange cultivar named after the famed oranges in València, Spain. It was first hybridized by pioneer American agronomist and land developer William Wolfskill in the mid-19th century on his farm in Santa Ana, southern California, United States, North America.
A seedless fruit is a fruit developed to possess no mature seeds. Since eating seedless fruits is generally easier and more convenient, they are considered commercially valuable.
Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from either parent. Asexual reproduction produces new individuals without the fusion of gametes, resulting in clonal plants that are genetically identical to the parent plant and each other, unless mutations occur.
Callose is a plant polysaccharide. Its production is due to the glucan synthase-like gene (GLS) in various places within a plant. It is produced to act as a temporary cell wall in response to stimuli such as stress or damage. Callose is composed of glucose residues linked together through β-1,3-linkages, and is termed a β-glucan. It is thought to be manufactured at the cell wall by callose synthases and is degraded by β-1,3-glucanases. Callose is very important for the permeability of plasmodesmata (Pd) in plants; the plant's permeability is regulated by plasmodesmata callose (PDC). PDC is made by callose synthases and broken down by β-1,3-glucanases (BGs). The amount of callose that is built up at the plasmodesmatal neck, which is brought about by the interference of callose synthases (CalSs) and β-1,3-glucanases, determines the conductivity of the plasmodesmata.
Polyembryony is the phenomenon of two or more embryos developing from a single fertilized egg. Due to the embryos resulting from the same egg, the embryos are identical to one another, but are genetically diverse from the parents. The genetic difference between the offspring and the parents, but the similarity among siblings, are significant distinctions between polyembryony and the process of budding and typical sexual reproduction. Polyembryony can occur in humans, resulting in identical twins, though the process is random and at a low frequency. Polyembryony occurs regularly in many species of vertebrates, invertebrates, and plants.
Gametogamy is sexual fusion – copulation or fertlization – of two single-celled gametes of different sex and the union of their gamete nuclei giving the zygote nucleus, as well as whole zygotic content.