Double fertilization

Double fertilization or double fertilisation (see spelling

differences) 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 (unless a type of self-incompatibility that The parts of a flower
acts in the stigma occurs in that particular species and is
activated), 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 mature embryonic sac of an unfertilized ovule is 7-cellular

and 8-nucleate. It is arranged in the form of 3+1+3 (from top to
bottom) i.e. 3 antipodal cells, 1 central cell (binucleate), 2
synergids & 1 egg cell. One sperm fertilizes the egg cell and the
other sperm fuses with the two polar nuclei of the large central
Double fertilization
cell of the megagametophyte. The haploid sperm and haploid egg
fuse to form a diploid zygote, the process being called syngamy,
while the other sperm and the diploid central cell fuse to form a triploid primary endosperm cell (triple
fusion). Some plants may form polyploid nuclei. The large cell of the gametophyte will then develop into
the endosperm, a nutrient-rich tissue which nourishes the developing embryo. The ovary, surrounding the
ovules, develops into the fruit, which protects the seeds and may function to disperse them.[1]

The two central cell maternal nuclei (polar nuclei) that contribute to the endosperm, arise by mitosis from
the same single meiotic product that gave rise to the egg. The maternal contribution to the genetic
constitution of the triploid endosperm is double that of the sperm.

In a study conducted in 2008 of the plant Arabidopsis thaliana, the migration of male nuclei inside the
female gamete, in fusion with the female nuclei, has been documented for the first time using in vivo
imaging. Some of the genes involved in the migration and fusion process have also been determined.[2]

Evidence of double fertilization in Gnetales, which are non-flowering seed plants, has been reported.[3]
Brief history
Double fertilization was discovered more than a century ago by Sergei Nawaschin in Kyiv,[4] and Léon
Guignard in France. Each made the discovery independently of the other.[5] Lilium martagon and
Fritillaria tenella were used in the first observations of double fertilization, which were made using the
classical light microscope. Due to the limitations of the light microscope, there were many unanswered
questions regarding the process of double fertilization. However, with the development of the electron
microscope, many of the questions were answered. Most notably, the observations made by the group of
W. Jensen showed that the male gametes did not have any cell walls and that the plasma membrane of the
gametes is close to the plasma membrane of the cell that surrounds them inside the pollen grain.[6]

Double fertilization in gymnosperms

A far more rudimentary form of double fertilization occurs in the sexual reproduction of an order of
gymnosperms commonly known as Gnetales.[3] Specifically, this event has been documented in both
Ephedra and Gnetum, a subset of gnetophytes.[7] In Ephedra nevadensis, a single binucleate sperm cell is
deposited into the egg cell. Following the initial fertilization event, the second sperm nucleus is diverted
to fertilize an additional egg nucleus found in the egg cytoplasm. In most other seed plants, this second
'ventral canal nucleus' is normally found to be functionally useless.[8] In Gnetum gnemon, numerous free
egg nuclei exist in female cytoplasm inside the female gametophyte. Succeeding the penetration of the
mature female gametophyte by the pollen tube, female cytoplasm and free nuclei move to surround the
pollen tube. Released from the binucleate sperm cell are two sperm nuclei which then fuse with free egg
nuclei to produce two viable zygotes, a homologous characteristic between families Ephedra and
Gnetum.[9] In both families, the second fertilization event produces an additional diploid embryo. This
supernumerary embryo is later aborted, leading to the synthesis of only one mature embryo.[10] The
additional fertilization product in Ephedra does not nourish the primary embryo, as the female
gametophyte is responsible for nutrient provision.[9] The more primitive process of double fertilization in
gymnosperms results in two diploid nuclei enclosed in the same egg cell. This differs from the
angiosperm condition, which results in the separation of the egg cell and endosperm.[11] Comparative
molecular research on the genome of G. gnemon has revealed that gnetophytes are more closely related to
conifers than they are to angiosperms.[12][13][14] The rejection of the anthophyte hypothesis, which
identifies gnetales and angiosperms are sister taxa, leads to speculation that the process of double
fertilization is a product of convergent evolution and arose independently among gnetophytes and
angiosperms.[15]

In vitro double fertilization

In vitro double fertilization is often used to study the molecular interactions as well as other aspects of
gamete fusion in flowering plants. One of the major obstacles in developing an in vitro double
fertilization between male and female gametes is the confinement of the sperm in the pollen tube and the
egg in the embryonic sac. A controlled fusion of the egg and sperm has already been achieved with poppy
plants.[16] Pollen germination, pollen tube entry, and double fertilization processes have all been observed
to proceed normally. In fact, this technique has already been used to obtain seeds in various flowering
plants and was named “test-tube fertilization”.[17]

Related structures and functions

Megagametophyte
The female gametophyte, the megagametophyte, that participates in double fertilization in angiosperms
which is haploid is called the embryonic sac. This develops within an ovule, enclosed by the ovary at the
base of a carpel. Surrounding the megagametophyte are (one or) two integuments, which form an opening
called the micropyle. The megagametophyte, which is usually haploid, originates from the (usually
diploid) megaspore mother cell, also called the megasporocyte. The next sequence of events varies,
depending on the particular species, but in most species, the following events occur. The megasporocyte
undergoes meiosis, producing four haploid megaspores. Only one of the four resulting megaspores
survives. This megaspore undergoes three rounds of mitosis, resulting in seven cells with eight haploid
nuclei (the central cell has two nuclei, called the polar nuclei). The lower end of the embryonic sac
consists of the haploid egg cell positioned in the middle of two other haploid cells, called synergids. The
synergids function in the attraction and guidance of the pollen tube to the megagametophyte through the
micropyle. At the upper end of the megagametophyte are three antipodal cells.

Microgametophyte
The male gametophytes, or microgametophytes, that participate in double fertilization are contained
within pollen grains. They develop within the microsporangia, or pollen sacs, of the anthers on the
stamens. Each microsporangium contains diploid microspore mother cells, or microsporocytes. Each
microsporocyte undergoes meiosis, forming four haploid microspores, each of which can eventually
develop into a pollen grain. A microspore undergoes mitosis and cytokinesis in order to produce two
separate cells, the generative cell and the tube cell. These two cells in addition to the spore wall make up
an immature pollen grain. As the male gametophyte matures, the generative cell passes into the tube cell,
and the generative cell undergoes mitosis, producing two sperm cells. Once the pollen grain has matured,
the anthers break open, releasing the pollen. The pollen is carried to the pistil of another flower, by wind
or animal pollinators, and deposited on the stigma. As the pollen grain germinates, the tube cell produces
the pollen tube, which elongates and extends down the long style of the carpel and into the ovary, where
its sperm cells are released in the megagametophyte. Double fertilization proceeds from here.[18]

See also
Fertilisation
Gamete
Megaspore
Oosphere
Ovule
Pollen
References
1. Berger, F. (January 2008). "Double-fertilization, from myths to reality". Sexual Plant
Reproduction. 21 (1): 3–5. doi:10.1007/s00497-007-0066-4 ([Link]
497-007-0066-4). S2CID 8928640 ([Link]
2. Berger, F.; Hamamura, Y. & Ingouff, M. & Higashiyama, T. (August 2008). "Double
fertilization – Caught In The Act". Trends in Plant Science. 13 (8): 437–443.
doi:10.1016/[Link].2008.05.011 ([Link]
PMID 18650119 ([Link]
3. V. Raghavan (September 2003). "Some reflections on double fertilization, from its discovery
to the present". New Phytologist. 159 (3): 565–583. doi:10.1046/j.1469-8137.2003.00846.x
([Link] PMID 33873607 ([Link]
[Link]/33873607).
4. Kordium EL (2008). "[Double fertilization in flowering plants: 1898-20&&&--". Tsitol. Genet.
(in Russian). 42 (3): 12–26. PMID 18822860 ([Link]
5. Jensen, W. A. (February 1998). "Double Fertilization: A Personal View". Sexual Plant
Reproduction. 11 (1): 1–5. doi:10.1007/s004970050113 ([Link]
0050113). S2CID 33416360 ([Link]
6. Dumas, C. & Rogowsky, P. (August 2008). "Fertilization and Early Seed Formation" ([Link]
[Link]/biologies/articles/10.1016/[Link].2008.07.013/).
Comptes Rendus Biologies. 331 (10): 715–725. doi:10.1016/[Link].2008.07.013 ([Link]
rg/10.1016%[Link].2008.07.013). PMID 18926485 ([Link]
485).
7. Carmichael, J. S.; Friedman, W. E. (1995-12-01). "Double Fertilization in Gnetum gnemon:
The Relationship between the Cell Cycle and Sexual Reproduction" ([Link]
g/content/7/12/1975). The Plant Cell. 7 (12): 1975–1988. doi:10.1105/tpc.7.12.1975 (https://
[Link]/10.1105%2Ftpc.7.12.1975). ISSN 1040-4651 ([Link]
4651). PMC 161055 ([Link]
PMID 12242365 ([Link]
8. Friedman, William E. (1990). "Sexual Reproduction in Ephedra nevadensis (Ephedraceae):
Further Evidence of Double Fertilization in a Nonflowering Seed Plant". American Journal of
Botany. 77 (12): 1582–1598. doi:10.1002/j.1537-2197.1990.tb11399.x ([Link]
02%2Fj.1537-2197.1990.tb11399.x). JSTOR 2444491 ([Link]
1).
9. Carmichael, Jeffrey S.; Friedman, William E. (1996). "Double Fertilization in Gnetum
gnemon (Gnetaceae): Its Bearing on the Evolution of Sexual Reproduction within the
Gnetales and the Anthophyte Clade". American Journal of Botany. 83 (6): 767–780.
doi:10.1002/j.1537-2197.1996.tb12766.x ([Link]
766.x). JSTOR 2445854 ([Link]
10. Friedman, W. E. (1995-04-25). "Organismal duplication, inclusive fitness theory, and
altruism: understanding the evolution of endosperm and the angiosperm reproductive
syndrome" ([Link] Proceedings of the
National Academy of Sciences. 92 (9): 3913–3917. Bibcode:1995PNAS...92.3913F (https://
[Link]/abs/1995PNAS...92.3913F). doi:10.1073/pnas.92.9.3913 ([Link]
org/10.1073%2Fpnas.92.9.3913). ISSN 0027-8424 ([Link]
424). PMC 42072 ([Link] PMID 11607532
([Link]
11. Friedman, William E. (1994). "The Evolution of Embryogeny in Seed Plants and the
Developmental Origin and Early History of Endosperm". American Journal of Botany. 81
(11): 1468–1486. doi:10.1002/j.1537-2197.1994.tb15633.x ([Link]
7-2197.1994.tb15633.x). JSTOR 2445320 ([Link]
12. Bowe, L. Michelle; Coat, Gwénaële; dePamphilis, Claude W. (2000-04-11). "Phylogeny of
seed plants based on all three genomic compartments: Extant gymnosperms are
monophyletic and Gnetales' closest relatives are conifers" ([Link]
c/articles/PMC18159). Proceedings of the National Academy of Sciences. 97 (8): 4092–
4097. Bibcode:2000PNAS...97.4092B ([Link]
92B). doi:10.1073/pnas.97.8.4092 ([Link] ISSN 0027-
8424 ([Link] PMC 18159 ([Link]
v/pmc/articles/PMC18159). PMID 10760278 ([Link]
13. Winter, Kai-Uwe; Becker, Annette; Münster, Thomas; Kim, Jan T.; Saedler, Heinz; Theissen,
Günter (1999-06-22). "MADS-box genes reveal that gnetophytes are more closely related to
conifers than to flowering plants" ([Link]
Proceedings of the National Academy of Sciences. 96 (13): 7342–7347.
Bibcode:1999PNAS...96.7342W ([Link]
W). doi:10.1073/pnas.96.13.7342 ([Link]
ISSN 0027-8424 ([Link] PMC 22087 ([Link]
[Link]/pmc/articles/PMC22087). PMID 10377416 ([Link]
377416).
14. Magallon, S.; Sanderson, M. J. (2002-12-01). "Relationships among seed plants inferred
from highly conserved genes: sorting conflicting phylogenetic signals among ancient
lineages" ([Link] American Journal of Botany. 89 (12):
1991–2006. doi:10.3732/ajb.89.12.1991 ([Link]
ISSN 1537-2197 ([Link] PMID 21665628 ([Link]
[Link]/21665628).
15. Chaw, Shu-Miaw; Parkinson, Christopher L.; Cheng, Yuchang; Vincent, Thomas M.; Palmer,
Jeffrey D. (2000-04-11). "Seed plant phylogeny inferred from all three plant genomes:
Monophyly of extant gymnosperms and origin of Gnetales from conifers" ([Link]
[Link]/pmc/articles/PMC18157). Proceedings of the National Academy of Sciences. 97
(8): 4086–4091. Bibcode:2000PNAS...97.4086C ([Link]
AS...97.4086C). doi:10.1073/pnas.97.8.4086 ([Link]
ISSN 0027-8424 ([Link] PMC 18157 ([Link]
[Link]/pmc/articles/PMC18157). PMID 10760277 ([Link]
760277).
16. Zenkteler, M. (1990). "In vitro fertilization and wide hybridization in higher plants". Crit Rev
Plant Sci. 9 (3): 267–279. doi:10.1080/07352689009382290 ([Link]
52689009382290).
17. Raghavan, V. (2005). Double fertilization: embryo and endosperm development in flowering
plants (illustrated ed.). Birkhäuser. pp. 17–19. ISBN 978-3-540-27791-0.
18. Campbell N.A.; Reece J.B. (2005). Biology (7 ed.). San Francisco, CA: Pearson Education,
Inc. pp. 774–777. ISBN 978-0-8053-7171-0.

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