Cotyledon

A cotyledon (/ˌkɒtɪˈliːdən/; "seed leaf"

from Latin cotyledon,[1] from Greek:
κοτυληδών kotylēdōn, gen.:
κοτυληδόνος kotylēdonos, from κοτύλη
kotýlē "cup, bowl") is a significant part of
the embryo within the seed of a plant,
and is defined as "the embryonic leaf in
seed-bearing plants, one or more of
which are the first to appear from a
germinating seed."[2] The number of
cotyledons present is one characteristic
used by botanists to classify the
flowering plants (angiosperms). Species
with one cotyledon are called
monocotyledonous ("monocots"). Plants
with two embryonic leaves are termed
dicotyledonous ("dicots").

Cotyledon from a Judas-tree (Cercis siliquastrum)

seedling.
Comparison of a monocot and dicot sprouting. The
visible part of the monocot plant (left) is actually
the first true leaf produced from the meristem; the
cotyledon itself remains within the seed

Schematic of epigeal vs hypogeal germination

Peanut seeds split in half showing the embryos with
cotyledons and primordial root.

Two week old Douglas fir with seven cotyledons.

Mimosa pudica seedling with two cotyledons and
the first "true" leaf with six leaflets.

In the case of dicot seedlings whose

cotyledons are photosynthetic, the
cotyledons are functionally similar to
leaves. However, true leaves and
cotyledons are developmentally distinct.
Cotyledons are formed during
embryogenesis, along with the root and
shoot meristems, and are therefore
present in the seed prior to germination.
True leaves, however, are formed post-
embryonically (i.e. after germination)
from the shoot apical meristem, which is
responsible for generating subsequent
aerial portions of the plant.

The cotyledon of grasses and many other

monocotyledons is a highly modified leaf
composed of a scutellum and a
coleoptile. The scutellum is a tissue
within the seed that is specialized to
absorb stored food from the adjacent
endosperm. The coleoptile is a protective
cap that covers the plumule (precursor to
the stem and leaves of the plant).

Gymnosperm seedlings also have

cotyledons, and these are often variable
in number (multicotyledonous), with from
2 to 24 cotyledons forming a whorl at the
top of the hypocotyl (the embryonic
stem) surrounding the plumule. Within
each species, there is often still some
variation in cotyledon numbers, e.g.
Monterey pine (Pinus radiata) seedlings
have 5–9, and Jeffrey pine (Pinus jeffreyi)
7–13 (Mirov 1967), but other species are
more fixed, with e.g. Mediterranean
cypress always having just two
cotyledons. The highest number reported
is for big-cone pinyon (Pinus
maximartinezii), with 24 (Farjon & Styles
1997).
The cotyledons may be ephemeral,
lasting only days after emergence, or
persistent, enduring at least a year on the
plant. The cotyledons contain (or in the
case of gymnosperms and
monocotyledons, have access to) the
stored food reserves of the seed. As
these reserves are used up, the
cotyledons may turn green and begin
photosynthesis, or may wither as the first
true leaves take over food production for
the seedling.[3]

Epigeal versus hypogeal

development
Cotyledons may be either epigeal,
expanding on the germination of the
seed, throwing off the seed shell, rising
above the ground, and perhaps becoming
photosynthetic; or hypogeal, not
expanding, remaining below ground and
not becoming photosynthetic. The latter
is typically the case where the cotyledons
act as a storage organ, as in many nuts
and acorns.

Hypogeal plants have (on average)

significantly larger seeds than epigeal
ones. They are also capable of surviving
if the seedling is clipped off, as meristem
buds remain underground (with epigeal
plants, the meristem is clipped off if the
seedling is grazed). The tradeoff is
whether the plant should produce a large
number of small seeds, or a smaller
number of seeds which are more likely to
survive.[4][5]

The ultimate development of the epigeal

habit is represented by a few plants,
mostly in the family Gesneriaceae in
which the cotyledon persists for a
lifetime. Such a plant is Streptocarpus
wendlandii of South Africa in which one
cotyledon grows to be up to 75
centimeters (2.5 feet) in length and up to
61 cm (two feet) in width (the largest
cotyledon of any dicot, and exceeded
only by Lodoicea). Adventitious flower
clusters form along the midrib of the
cotyledon.[6] The second cotyledon is
much smaller and ephemeral.

Related plants may show a mixture of

hypogeal and epigeal development, even
within the same plant family. Groups
which contain both hypogeal and epigeal
species include, for example, the
Southern Hemisphere conifer family
Araucariaceae,[7] the pea family,
Fabaceae,[4] and the genus Lilium (see
Lily seed germination types). The
frequently garden grown common bean,
Phaseolus vulgaris, is epigeal, while the
closely related runner bean, Phaseolus
coccineus, is hypogeal.
History
The term cotyledon was coined by
Marcello Malpighi (1628–1694).[a] John
Ray was the first botanist to recognize
that some plants have two and others
only one, and eventually the first to
recognize the immense importance of
this fact to systematics, in Methodus
plantarum (1682).[3][11]

Theophrastus (3rd or 4th century BC) and

Albertus Magnus (13th century) may also
have recognized the distinction between
the dicotyledons and
monocotyledons.[12][13]
Notes
a. The Oxford English Dictionary
attributes it Linnaeus (1707–1778)
"1751 Linnaeus Philos. Bot. 54.
Cotyledon, corpus laterale seminis,
bibulum, caducum" [8] and 89, [9] by
analogy with a similar structure of
the same name in the placenta.[10]

References
1. Short & George 2013, p. 15 .
2. OED 2019.
3. Vines, Sydney Howard (1913),
"Robert Morison 1620—1683 and
John Ray 1627—1705", in Oliver,
Francis Wall (ed.), Makers of British
 botany, Cambridge University Press,
pp. 8–43
4. Charles R. Tischler; Justin D. Derner;
H. Wayne Polley; Hyrum B. Johnson,
Response of Seedlings of Two
Hypogeal Brush Species to CO2
Enrichment , Fort Collins, CO: U.S.
Department of Agriculture, Forest
Service, Rocky Mountain Research
Station, pp. 104–106
5. Baraloto, C.; Forget, P.-M. (2007),
"Seed size, seedling morphology, and
response to deep shade and damage
in neotropical rain forest trees",
American Journal of Botany, 94 (6):
 901–11, doi:10.3732/ajb.94.6.901 ,
PMID 21636459 , S2CID 24272337
6. Perry, Frances and Leslie Greenwood
(1972). Flowers of the World.
London: Hamlyn Publishing Group.
p. 47.
7. Hiroaki Setoguchi; Takeshi Asakawa
Osawa; Jean-Christophe Pintaud;
Tanguy Jaffré; Jean-Marie Veillon
(1998), "Phyloghuhenetic
relationships within Araucariaceae
based on rbcL gene sequences" ,
American Journal of Botany, 85 (11):
1507–1516, doi:10.2307/2446478 ,
JSTOR 2446478 , PMID 21680310
8. Linnaeus 1751, p. 54 .
 9. Linnaeus 1751, p. 89 .
10. OED 2015.
11. Greene, E. L. & Egerton, F. N. (ed.)
(1983). Landmarks of Botanical
History: Part 2. Stanford: Stanford
University Press, p. 1019, note 15,
[1] .
12. "Bioetymology: Origin in Biomedical
Terms: cotyledon, monocotyledon
(plural usually monocots),
dicotyledons(plural usually dicot)" .
bioetymology.blogspot.com.br.
Retrieved 6 April 2018.
13. Greene, E. L. & Egerton, F. N. (ed.)
(1983), p. 1019, note 15.

Bibliography
Linnaeus, Carl (1755) [1751]. Philosophia
botanica : in qua explicantur fundamenta
botanica cum definitionibus partium,
exemplis terminorum, observationibus
rariorum, adiectis figuris aeneis . originally
published simultaneously by R. Kiesewetter
(Stockholm) and Z. Chatelain (Amsterdam).
Vienna: Joannis Thomae Trattner. Retrieved
13 December 2015.
Mirov, N. T. (1967). The Genus Pinus.
Ronald Press Company, New York.
Farjon, A. & Styles, B. T. (1997). Pinus
(Pinaceae). Flora Neotropica Monograph 75:
221-224.
"Cotyledon" . Oxford English Dictionary
(Online ed.). Oxford University
Press. (Subscription or participating institution
membership required.)
 Short, Emma; George, Alex (2013). A primer
of botanical Latin with vocabulary . New
York: Cambridge University Press.
ISBN 9781107693753. Retrieved
14 December 2015.

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