Cell Development During Early Embryogenesis in Capsella and Gossypium
Edward G, Pollock; William A. Jensen
American Journal of Botany, Vol. 51, No. 9. (Oct., 1964), pp. 915-921.
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‘Thu Jul 13 19:02:25 2006‘Amer. Jour. Bot. 51(0): 915-021. 1964,
CELL DEVELOPMENT DURING EARLY EMBRYOGENESIS IN
CAPSELLA AND GOSSYPIUM!
Epwarp G. Poutock aNp Wittiam A. JENSEN
Department of Biology, San Fernando Valley State Collage, Northridge, California
and
Department of Botany, University of California, Berkeley, California
ABSTRACT
‘The early stages of embryo development in Goeeypium hireutum (cotton) and Copslia bursa-
pastoris were examined with rgard to patterns of eell development, embryo and cell size, and dis-
tribution of eel di
following the first di
more divis
sine was larger than the sygote. Distnetive pattern of eal di
eat that changes in groups of cells undergoing mitosis are of fundamental importance in
understanding the development of form in the embryo. A greater
and
12. A striking reduction inthe total sie of the cotton embryo was observed
in of the embryo, This decreas in total embryo size continued for several
ns, and it was not until the embryo contained approximately 75 cells that its total
1 were fond in both embryos
gree of variation in develop-
ment of eel Hineages than is generally reported was oberved in both embry.
‘Te pevevorwmxr of the plant embryo has
Deen studied in detail for many years (Hanstein,
870; Soeges, 1914; Johansen, 1950; Maheshwari,
1950; Wardlaw, 1955). These studies have been
primarily morphological and desexiptive in nature.
‘The need for data on the biochemical changes
associated with plant embryogenesis is increasing
as the problem of plant cell development is ap-
‘proached in terms of the embryo, ‘This paper is
intended as the morphological basis for an ex-
‘ended series of investigations which will examine
Various biochemical aspects of ‘embryogenesis,
‘With this consideration in mind, the analysis was
le as quantitative as possible, particularly wit
regard to cell number, eel size, and the distribu-
tion of cell divisions. ‘The result is that new data,
‘onthe development of 2. well-known” plant
embryos (Capsella bursa-pastoris and. Gossypium
drirautum) were obtained.
‘Marensats 4xp aztions—The plants used
inthis study were Capsella bursa-pastoris. L.
‘Medio, the shepherd's purse, and Gossypium
hrireutim L., cultivated ‘cotton. ‘The “Capella
plants were krovn from, std from the Botan
arden, University of California, Berkeley. ‘The
cotton plants were grown from seed of variety
‘M8048 ‘(a double haploid) developed by Dr.
James Meyers at the Delta. Branch Experimental
Station, Stoueville, Mississippi
Whole silicles of Capsela and excised ovules of
Gossypivm vere chemically fixed, ‘The lateral
‘Portions of the silieles were cut off to allow for
better penetration of the fixative, For the same
reason, the chalazal ends of ovules older than 5
+ Received for publication April 14, 1968
[he Jounsas. for September (51: 809-013) wae ianued
‘Aatemteax Jownsas or Boras, Vol. 1! No.
days after
were fixed
fluid Jensen, 1962), FPA (formalin, propionic
acid, and ethyl algohol), and Carnoy’s. fluid
Gohansen, 1940). All chemically fixed material
was embedded in’ paraffin via the standard TBA
series, The tissues were sectioned at 5u and stained
‘with Heidenhain’s iron hematoxylin; then counter
stained with Fast Green or Orange G (lohansen,
1940).
Celi_number—The average cell number was
determined for each stage in embryo development.
by counting the miclei of the cells in serial sections
of embryos. Six embryos were counted for each of
the stages of embryo growth described in the pre-
ceding section,
Dictrution of mit.tie fgures—The number of
cells undergoing division’ were counted in serial
sections of entire embryos. All nuclei showing
late-prophase to early-telophase figures were
counted. The various stages in embryo develop-
ment, beginning with the initial cleavages of the
zygote up to and including the torpedo stage,
‘were exainined. Counts were made on 4 embryos
at each stage of growth in Gossypium, and
embryos were counted for each stage of growth in
Capsella, The counts were totalled and plotted on
outline drawings of ‘the succeading, stags of
‘Bnrjo ize—With the ad ofa eamera hid,
outline drawings were made of median longitudi=
nal sections of embryos at all stages of growth. up
to and including the torpedo stage. ‘The areas of
the outline drawings were measured with a plani-
meter; the readings were taken directly off the
planimeter scale in square millimeters and con-
tember 90, 1964
1964
1516
U9¢@
" 12 13
Fig. 1-18. Stages of early cleavage in Gostypium.—Fig,
1. Eeg.—Fig, 2. The nygote—Fig. 3. The 2eelled embryo,
Fig. 47. Threceslled embryox—Fig. 8—A t-eclled
embryo—Fig. 9-12. Subsequent staget in development,
showing the variable cleavage patterns Fig, 18, An early
slobular embryo showing the differentiation of the derma-
{ogen. All figures X805.
verted to square-micron units to obtain the real
areas of the sections measured. The area of the
‘median longitudinal section was assumed to be a
valid index of embryo growth as the embryo i
AMERICAN JOURNAL OF nOTANY
[Vol 51
essentially isodiametric up to the early heart
stage; and the subsequent increase in embryo size
is essentially Tinear up to and including the torpedo
stage, Fight embryos were drawn and measured.
in this manner for each of the stages of embryo
growth.
Call size—The same outline drawings prepared
to measure embryo size were used to determine cell
size. The number of cells in each median longitu-
inal section was counted. ‘The average area of the
cells was then determined by dividing the area of
the same section by the coll number of that
section.
Rusvurs—Morphological analysis—Hlistogenic
atterns—The cleavage patterns of Capeella are
well known (Johansen, 1950; Wardlaw, 1955). Up
to the formation of the first'cells of the dermato-
gen, the sequence of cleavage is apparently always
the'same (Sooges, 1914; 1919). Beyond this point
in histogenesis, it was observed that the cells
arising from subsequent divisions were not always
disposed in exactly the same manner in each
embryo. These differences in the displacement, of
cells for different. embryos were the exception
rather than the rule. For example, the 8 cells of
the young dermatogen occasionally divided anti-
elinally and synchronously, producing a 10-celled
dermatogen, while the 8 central eells were as yet
ided. A more frequent oceurrence was the
longitudinal division of each of the oetant cells
forming 16 central cells before any of the 8 cells
of the dermatogen divided
Other deviations from a precise cleavage pat
tern were observed in following the cleavages in
the young globular stage to the early heart stage,
Normally, the upper tier of 4 eolls in the octant
stage gives rise to the shoot apex and cotyledons,
while the cells of the lower tier give rise to the
hypocotyl (Johansen, 1950; Maheshwari, 1950;
Wardlaw, 1955), but. this’ need not be true.
‘Though synchrony of cell division exists up to the
cctant stage with its firstformed cells of the
dermatogen, irregularities in the sequence of divi-
ions soon become apparent with subsequent,
divisions. In following embryo development from
the youngest globular stage up to the early heart
stage, it was observed that more divisions were
‘occurring in the upper half of the sphere than in
longitudinal sections through developing ovules of Gossypium showing progressive development of
4, The large zygote (2) attached to the nu
lar tise (nt) of the ovule, X