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Figure 3.3: Different types of ovules: A. Orthotropous ovule B. Anatropous ovule C.

Amphitropous ovule D. Campylotropous ovule E. Hemianatropous ovule F. Circinotropous ovule

Considering the extent of development of the nucellus and position of sporogenous

cell, ovule can also be categorized as

1. Tenuinucellate type
2. Crassinucellate type

1. Tenuinucellate type:
The archesporial cell in nucellus is hypodermal in origin. When it directly functions as the
megaspore mother cell, the position of this megaspore mother cell (sporogenous cell) is
also hypodermal. So the nucellar tissue around it also remains single-layered. Such ovules,
where the position of sporogenous cell is hypodermal and the nucellar tissue around it
remains single-layered, are called tenuinucellate (Figure 3.4) type of ovule.

Figure 3.4: A. Tenuinucellate type of ovule in Orchis maculates, in which the megasporocyte
occurs directly below the nucellar epidermis; B.Crassinucellate type of ovule in Quisqualis indica
(note deeply embedded position of megasporocyte and the active periclinal divisions in the nucellar
epidermis). [Adapted from An Introduction to the Embryology of Angiosperms by P. Maheshwari.
N.Y.: McGraw-Hill Book Company, Inc., 1950]

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2. Crassinucellate type:
Unlike tenuinucellate type, the hypodermal archesporial cell divides into two cells by
means of transverse division. The outer one is parietal cell and an inner one is sporogenous
cell. The parietal cell may either remain undivided or undergo a few divisions. These
divisions are both periclinal as well as anticlinal. As a result of this the sporogenous cell
becomes embedded in the massive parietal tissue. The sporogenous cell may also be
embedded in the massive nucellus by divisions in the nucellar epidermis instead of division
in archesporial cell. All such ovules where the sporogenous cell becomes sub-hypodermal,
by either above two means, are called crassinucellate (Figure 3.4).

Tenuinucellate ovule: where the sporogenous cell is hypodermal and the

nucellar tissue around it remains single layered

Crassinucellate ovule: where the sporogenous cell becomes sub-hypodermal,

3.4 DEVELOPMENT OF THE FEMALE GAMETOPHYTE
either due to formation of parietal cells, or due to divisions in the nucellar
epidermis, or both
In general female gametophyte development occurs within the ovule (megasporangium)
and completes in two phases:

1. Megasporogenesis
2. Megagametogenesis
Development of the megaspore from megaspore mother cell is known as
megasporogenesis and a process of cell proliferation and differentiation in megaspore to
develop into multicellular female gametophyte is known as megagametogenesis.

Angiosperm plants follow many different patterns of female gametophyte development.

The common developmental pattern exhibited by over 70% of flowering plants is referred
to as the Polygonum-type. Example- Poaceae (e.g., maize, rice, wheat), Phaseoleae (e.g.,
beans, soyabean), Brassicaceae (e.g., Brassica), Malvaceae (e.g., cotton), and Solanaceae
(e.g., pepper, tobacco, tomato, potato, petunia) and most apomictic species (Maheshwari
1950, Willemse and van Went 1984, Huang and Russell 1992). It is known as
Polygonum-type because it was first described in Polygonum divaricatum (Strasburger,
1879; Maheshwari, 1950).

Megasporogenesis

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“Development of the megaspore within the ovule (megasporangium) from megaspore

mother cell is known as megasporogenesis.”

At the micropylar end, one of the hypodermal cells in the nucellus differentiates and
functions as the archesporial cell (archesporium). Being large in size, having dense
cytoplasm and large nucleus, it becomes more prominent than its surrounding cells.
Therefore is easily distinguishable from the other cells.

Depending on the basis of position of sporogenous cell, ovule can be categorized into two
types-1.tenuinucellate type and 2.crassinucellate type (described earlier).

So here in tenuinucellate type of ovule, the archesporial cell directly functions as

megaspore mother cell (MMC) and in crassinucellate type of ovule the archesporial cell
do not directly behave as MMC and instead of that it divides periclinally into two cells.
An outer primary parietal cell (towards epidermis) and an inner primary sporogenous cell.
Now this primary sporogenous cell functions as the megaspore mother cell.

Megaspore mother cell, also known as megasporocyte is diploid in nature, representing

the last cell of sporophytic generation. It undergoes meiosis i.e. reduction division. As a
result of this four haploid megaspores (tetrad) are formed. Tetrad of megaspores may be
arranged in linear, T-shape, isobilateral or tetrahedral patterns. T-shaped tetrad arises due
to vertical division in the micropylar dyad cell and transverse division in the chalazal dyad
cell. In case of linear arrangement which is most common, after the first meiotic division,
the wall is laid down transversely, forming a dyad. The second meiotic division in the two
dyad cells is also transverse. In this way a row of four haploid megaspore cells (linear
tetrad) is formed (Figure 3.5 and 3.6).

Figure 3.5-Formation of 4 haploid megaspores from diploid megaspore mother cell

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Figure 3.6. Megasporogenesis in the ovule of Hydrilla verticillata. A. hypodermal sporangial

initial. B. Periclinal division of sporangial initial into parietal cell and primary sporogenous cell C.
results of anticlinal division of parietal cell D. megasporocyte situated below the layer of wall cells
E-F. formation of dyad cells G. linear tetrad of megaspores (Adapted from An Introduction to the
Embryology of Angiosperms by P. Maheshwari. N.Y.: McGraw-Hill Book Company, Inc., 1950)

In the linear tetrad, the lowermost megaspore (the chalazal megaspore) enlarges and
becomes functional, while rest three megaspores of tetrad do not participate in the
formation of female gametophyte and degenerate.

The functional megaspore now forms the female gametophyte (embryo sac).

If you remember the microsporogenesis where a haploid microspore was said to be the first
cell of male gametophyte, similarly here megaspore is known as the first cell of female
gametophyte. In angiosperms the development of the female gametophyte is completely
endosporous means within the megaspore.

Megagametogenesis

Megagametogenesis could be defined as a process of cell proliferation and differentiation

in megaspore cell to develop into multicellular female gametophyte.

Initially, the surviving megaspore undergoes free nuclear divisions (without cytokinesis)
resulting into a multinucleate coenocyte. Subsequently, development of cell walls around
these nuclei leads to the development of a female gametophyte (Figure 3.5).

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In case of Polygonum-type of female gametophyte, the single functional (surviving)

megaspore increases in size. After enlargement it undergoes two rounds of mitosis without
cytokinesis. As a result of which a four-nucleate coenocyte formation takes place having
two nuclei at each pole and all these nuclei divide again third time. After third mitosis
eight-nucleate coenocyte formation takes place having four nuclei at each pole.

Now one nucleus from each pole migrates toward the center of the developing female
gametophyte known as the polar nuclei which may or may not fuse together either before or
after the entry of pollen tube.

Ultimately an eight nucleated structure consisting of three antipodal cells, two polar nuclei
(central cell), two synergid cells and one egg cell is formed.

The central cell having two identical haploid nuclei and is therefore called as homodiploid.
The other all cells inherit single haploid nuclei (Figure 3.7).

Figure 3.7- Development of female gametophyte of normal type

Figure 3.8 and Figure 3.9 showing megasporogenesis (the diploid megaspore mother cell
undergoes meiosis and gives rise to haploid megaspores) and megagametogenesis (one of
the megaspores develops into the mature female gametophyte), respectively in
Arabidopsis.

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Figure 3.8-Megasporogenesis in Arabidopsis.

A. Apical region of a young, finger like ovule primordium. The megaspore mother cell forms from a
sub-epidermal cell at the distal end of the ovule primordium. L1 is the outer layer of cells.
B. Steps of megasporogenesis. Ovule promordia arise as finger-like projections from the placenta.
The megaspore mother cell undergoes meiosis and forms four megaspores. Three of the
megaspores undergo cell death. The chalazal-most megaspore survives, becomes the functional
megaspore and undergoes megagametogenesis.
Black circles/ovals represent nuclei.dm: degenerating megaspores; fm: functional megaspore; ii:
inner integument; L1: epidermal layer of the ovule promordium; mmc: megaspore mother cell; mt:
meiotic tetrad; oi: outer integument.
Source:Drews and Koltunow(2011)

Fig. 3.9-Megagametogenesis in Arabidopsis

A. Steps of megagametogenesis emphasizing development within the ovule.

B. Stages of megagametogenesis (Christensen et al. 1998). The megaspore contains a single
nucleus (stage FG1). The nucleus undergoes two rounds of mitosis, producing four-nucleate
coenocytes, with two nuclei at each pole separated by a large central vacuole (stage FG4). During
a third mitosis, phragmoplsta and cell plates form between sister and non-sister nuclei and the
nuclei become completely surrounded by cell walls (stage FG5). During cellularization, the polar
nuclei migrate towards the centre of the female gametophyte and fuse before fertilization. These
events produce a seven-celled structure consisting of three antipodal cells, one central cell, two

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synergids, and one egg cell. If the female gametophyte is not fertilized, the antipodal cells
eventually degenerate (stage FG7, not shown).
White areas represent vacuoles and black circles/ovals represent nuclei.
Ac: antipodal cells; cc: central cell; ccn: central cell nucleus; ch: chalazal region of the ovule; ec:
egg cell; f: funiculus; fg: female gametophyte; fm: functional megaspore; ii: inner integument; m:
megaspore; mp: micropyle; oi: outer integument; pn: polar nuclei; sc: synergid cells.
Source: Drews and Koltunow (2011)

Steps of development of the female gametophyte or embryo sac in detail:

The development of embryo sac begins as the functional megaspore elongates. You know
that in most cases the lowermost megaspore (chalazal) of the linear tetrad becomes
functional and rest three degenerates. The elongation is largely along the
micropylar-chalazal axis.

Micropylar- end

Chalazal- end

Functional megaspore Embryo sac

The first nuclear division (post- meiotic mitosis) in the megaspore is not followed by wall
formation. A large central vacuole appears between the two daughter nuclei. Initially there
is no vacuole in the cytoplasm of the megaspore but later small vacuoles appear which may
fuse to form large vacuole. As the vacuole expands, the nuclei pushed toward opposite
poles of the cell. In this way each pole has one nucleus. Now both the nuclei divide twice,
forming four nuclei at each pole. All the divisions are mitotic and without wall formation.

Dear students you can understand that at this stage all the eight nuclei are present in the
common cytoplasm because after haploid megaspore formation all the nuclear divisions are
not followed by cell wall formation. After the last nuclear division (when there are eight
nuclei) the cell undergoes appreciable elongation, so that it looks like sac. The picture must
be clear in your mind that out of these eight nuclei, four are at the micropylar-end and four
at the chalazal-end.

Finally the embryo sac becomes organized. Three nuclei at the micropylar-end of the
embryo sac organize into egg apparatus and the fourth one is left free in the cytoplasm of
the central cell and moves toward the centre as the upper polar nucleus. The egg apparatus

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(only upper portion) is attached to the wall of the embryo sac at the micropylar-end, its
major portion is surrounded by the middle cell. In the egg apparatus, the middle cell is egg
cell (round, also known as ovum or oosphere) and the rest two side cells are synergids
which are flask shaped. Only the synergid cells are in direct contact with the wall of the
embryo sac. The central egg cell is situated in such a way that its upper portion is slightly
below the apices of the synergids and seems hanging between and below them. All the
three cells are of same length therefore the egg cell extends a little more towards the centre
in comparison to the synergids. Out of the four nuclei at the chalazal-end of the embryo sac,
three nuclei forms three antipodal cells and the fourth one moves toward the centre as the
lower polar nucleus.

You have read that one nucleus from each pole moves to the center of the embryo sac, here
they may fuse forming the fusion or secondary nucleus. The secondary nucleus (if fusion
occurred between two polar nuclei) or two polar nuclei (if there is no fusion) remains at the
center.

The haploid nucleus of the megaspore divides mitotically (non-reductional division) which
organize in a definite manner within the embryo sac. Three nuclei at the micropylar-end, three
at the chalazal-end and the remaining two migrate to the centre of the embryo sac. Three nuclei
at the micropylar-end organize into an egg apparatus. The central large cell of the egg
apparatus is called egg cell (female gamete) which is partially surrounded by two lateral
synergid cells. Three nuclei of the chalazal-end form antipodal cells. The two nuclei which
migrates to the centre, called polar nuclei. These polar nuclei later fuse to form a single diploid
secondary nucleus (a central cell).
These events result in a mature seven celled structure called female gametophyte or
embryo sac consisting of three antipodal cells, one central cell having two polar nuclei,
two synergid cells, and one egg cell. Since, this type of embryo sac develops from a
single megaspore and has eight nuclei, it is said to be monosporic-8-nucleate embryo
sac or Polygonum -type of embryo sac. Throughout development, the female gametophyte
exhibits a polarity along its chalazal-micropylar axis.

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Important Points
- The female gametophyte (embryo sac) is 7-celled (mostly), 8-nucleate structure having
three cells of egg apparatus (two synergid cells and one egg cell) at the micropylar-end,
three cells (antipodal cells) at the chalazal-end and one cell (centre cell) in the centre having
two polar nuclei.
-This type of embryo sac is designated as the Polygonum-type, formed as a result of three
divisions in functional megaspore.

-This mode of embryo sac development occurs in the majority type of flowering plants.
According to Davis (1966), about 81 per cent of the families show Polygonum-type of
embryo sac development.

- Cells of the egg apparatus and the antipodal cells are uninucleate and haploids whereas the
central cell is binucleate or diploid.

The entire embryo sac is enclosed within diploid sporophytic tissues called integuments,
which will constitute the seed coat in the mature seed.
Detailed structure of the mature female gametophyte

Transmission electron microscopic structure of mature female gametophyte in Arabidopsis

(Figure 3.10) has been described (Mansfield et al., 1991; Murgia et al., 1993; Kasahara et
al., 2005; Kagi et al., 2011). The mature female gametophyte in Arabidopsis is ~105 µm
long and ~25 µm wide and is Polygonum-type.

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Chalazal-end

A: Ovule B: Female gametophyte

Fig. 3.10. A: The Arabidopsis Female gametophyte within the ovule. B: The mature female
gametophyte. The gray area represents cytoplasm, the white area represents vacuoles and the black
area represents the nuclei. ac: antipodal cells; cc: central cell; ch: chalazal region of the ovule; ec:
egg cell; f: funiculus; mp: micropyle; sc: synergid cell; sn: secondary nucleus. Picture courtesy of
Yadegari and Drews (2004).

The egg cell and central cell are polarized such that the nuclei of both cells lie very close to
Micropylar-end
each other (Drews and Koltunow 2011). This feature is very important during double
fertilization because these two nuclei are involved or we can say are the targets of the two
male nuclei.

The cell wall is either absent or discontinuous at the junction of egg, synergid and central
cell so that the plasma membrane of these cells is in direct contact with each other
(Mansfield et al., 1991; Kasahara et al., 2005). This discontinuation or absence of cell wall
in these regions provide direct access of the male gametes or sperm cells to the fertilization
targets i.e. egg nucleus in the egg cell and secondary nucleus in the central cell because one
of the synergid cells receives two male gametes from the growing pollen tube (Figure
3.11).

The synergid cell wall is further modified. At the micropylar site it is thickened and
extensively invaginated, forming a structure known as filiform apparatus. Formation of
filiform apparatus by means of invaginations in the cell wall of synergids helps in
increasing the surface area of the plasma membrane in this region. It also contains a high
concentration of secretory organelles. It is supposed that it may facilitate transport of
substances into and out of the synergid cells (Drews and Koltunow 2011) (Figure 3.11).
Findings of cytological staining properties in other plant species concluded that the filiform
apparatus is composed of cellulose, hemicellulose, pectin, callose, and proteins.

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Schematic development of female gametophyte, based on the model plant Arabidopsis is

given in Figure 3. 12.

Fig. 3.12 Schematic development of female gametophyte, based on the model plant Arabidopsis.

(A) The megaspore mother cell (MMC) is surrounded by epidermal cells of the nucellus (Nu) prior
to undergoing meiosis to generate four spores. At this stage, the formation of the outer (OIn) and
inner (IIn) integuments has just initiated.
(B) Diagram of MMC asymmetric meiosis that generates four spores (tetrad). Three of these
undergo programmed cell death. The proximal (chalazal) megaspore becomes the functiona
lmegaspore (FM). (C) FG1 stage. The FM is teardrop-shaped andundergoes the first mitotic
division.
(D) FG2 stage. The female gametophyte comprises two nuclei. The nucellus (Nu) is enclosed by the
OIn, but not the IIn integuments.
(E) Stages FG3 to FG7. The female gametophyte comprises two nuclei, separated by a large
vacuole (V), that undergo second and third mitotic divisions to generate the eight-nucleate mature
embryo sac at the FG5 stage. Subsequent cellularization (FG6 stage) results in the formation of
seven cells: two synergid cells (SC); one egg cell (EC); one central cell (CC) carrying two polar
nuclei (PN); and three antipodal cells (AC). By FG7, the two polar nuclei have fused to form the
central cell nucleus (CCN), and the antipodal cells degenerate.
CN, chalazal nucleus; CV, central vacuole; DM, degenerating megaspores; Fu, funiculus; IIn,
inner integuments; MMC, megaspore mother cell; MN, micropylar nucleus; Nu, nucellus; OIn,
outer integuments.
Source: Sundaresan, and Alandete-Saez. Development. 2010 137: 179-189; doi: 10.1242/dev.030346

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The egg cell The embryo

Two synergid cells For pollen tube attraction

The central cell The endosperm

Female gametophyte Polarity

Throughout the process of development, the female gametophyte exhibits a polarity along
its chalazal-micropylar axis. We can say that the ovule and female gametophyte are
polarized structures as shown in figure 1. The integuments form a pore called micropyle at
the anterior pole and its chalazal pole is the posterior end that joins the funiculus (inverted
position in anatropous ovule).

3.5 ALTERNATIVE PATTERNS OF FEMALE

GAMETOPHYTE DEVELOPMENT
The Polygonum-type of female gametophyte development is the most common pattern.
Many other patterns of development of female gametophyte also exist.
These different patterns are because of the variations in megasporogenesis and
megagametogenesis both.
3.5.1. Patterns due to variation in megasporogenesis
Megasporogenesis among angiosperms exhibit three main patterns, referred
to as monosporic, bisporic, and tetrasporic (Figure 3.13).
These three patterns differ mainly in - whether wall (cell plate) formation occurs after each
meiotic division or not.

This decides the number of meiotic products that contribute to the formation of the mature
female gametophyte, and also the pattern.

In the monosporic pattern, each phase of meiotic division is accompanied by wall

formation, resulting in four uninucleate megaspores (linear tetrad). Only one becomes
functional which forms the female gametophyte (embryo sac).

In the bisporic pattern, cell wall develops after meiosis I but not after meiosis II resulting
in two binucleate cells. One of the dyad cells degenerates, resulting in a single functional

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dyad cell possessing two megaspore nuclei. This functional dyad cell with two megaspore
nuclei contributes towards the formation of female gametophyte.

In the tetrasporic pattern, neither first nor second meiotic division is accompanied by
wall formation resulting in the formation of four megaspore nuclei inside the cell. All four
megaspore nuclei participate in the formation of female gametophyte.

Monosporic pattern- one

megaspore contributes to formation
of the mature female gametophyte.
Bisporic pattern - two megaspore
nuclei (dyad cell) contribute to
formation of the mature female
gametophyte.
Tetrasporic pattern – All the four-
megaspore
. nuclei contribute to
formation of the mature female
gametophyte.
Fig. 3.13- Patterns of megasporogenesis

3.5.2. Patterns due to variation in megagametogenesis.

Same general pattern as discussed above for Arabidopsis i.e. a phase of nuclear
proliferation without cytokinesis followed by cellularization and differentiation is reported
in majority of angiosperm species.

Variations arise because of:

-the number of nuclei within the megaspore that give rise to the female gametophyte
(i.e. the type of megasporogenesis)
-the number of mitoses prior to cellularization
-the timing of fusion of the polar nuclei
-whether or not additional mitoses occur after cellularization.
(Maheshwari 1950; Willemse and van Went 1984; Haig 1990; Huang and Russell
1992; Yadegari and Drews 2004).
As in maize, exhibiting Polygonum- type of female gametophyte, the polar nuclei do
not fuse until fertilization and the antipodal cells proliferate into 40 or more cells (Diboll
and Larson 1966, Diboll 1968).

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Therefore depending on the basis of involvement of number of megaspore nuclei

in its formation, the embryo sac can also be called as: Monosporic, Bisporic and
Tetrasporic

3.5.3 Types of Female gametophyte/Embryo Sac Development

Maheshwari (1950) has developed a classification of the various types and subtypes of
embryo sac development in the angiosperms. His classification is fundamentally based on:
(1) The number of spores or spore-nuclei which enter into the formation of embryo sac
(2) The total number of nuclear divisions which occur during megasporogenesis and
megagametogenesis and
(3) The number, arrangement and chromosome number of the nuclei in the mature embryo
sac.

1. Monosporic Type:
Here megasporogenesis results in four well-defined megaspores, one of which gives rise to
the embryo sac, rest three megaspores degenerate. Most commonly the megaspore farthest
from the micropyle is functional. In this type of embryo sac all the nuclei are genetically
identical because they are formed through mitosis of a single nucleus.
Monosporic embryo sacs are further divided into two types.

1. Polygonum type (8 nucleate)

2. Oenothera type (4 nucleate)
(i) Polygonum type (8 nucleate): As described earlier, it is formed by the chalazal
megaspore of the tetrad and is eight nucleate. The mature Polygonum type of embryo sac
comprises a 3-celled egg apparatus, three antipodal cells and a binucleate central cell
means seven -celled, eight- nucleate monosporic embryo sac (Figure 3.14).

Chalazal megaspore

Figure 3.14- Polygonum type embryo sac

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This type of embryo sac is the most common and is, therefore, commonly designated as the
"Normal type." However, it is also designated as the Polygonum- type because it was
reported for the first time in Polygonum divaricatum by Strasburger (1879).

(ii) Oenothera type (4 nucleate): Oenothera type of embryo sac is derived from the
functional micropylar megaspore of the tetrad and is four nucleate. The mature embryo sac
consists of an egg apparatus and a uninucleate central cell (polar nucleus). Oenothera type
of embryo sac is found in Onagraceae family.

Geert, in 1908, found that in Oenothera lamarckiana the embryo sac is usually formed
by the micropylar megaspore of the tetrad, which undergoes only two nuclear divisions
instead of the usual three occurring in the Polygonum type of embryo sac. In this way, 4
nuclei are produced which organize into the two synergids, the egg and a single polar
nucleus. Since the third division is omitted and all the nuclei are situated in the micropylar
part of the developing embryo sac, there is neither a lower polar nucleus nor any antipodal
cells (Figure 3.15).

Micropylar megaspore

Fig. 3.15-Oenothera type embryo sac

The Oenothera type is of particular morphological interest because following double

fertilization, the primary endosperm nucleus is diploid rather than triploid or polyploid as
in other angiosperms.

2. Bisporic Type:

As described earlier, in this type of embryo sac as the name indicates two megaspore
nuclei participate in its formation. After first meiotic division a dyad is formed by wall
formation. Only one of the dyad cells undergoes the second meiotic division and the other
one degenerates. In the functional dyad, cell division is not followed by wall formation and
so both the megaspore nuclei (haploid) participate in the formation of the embryo sac. Each
megaspore nucleus undergoes two mitotic divisions forming eight nuclei and mature
embryo sac has the same organization like that of the Polygonum- type.

So you can say the bisporic embryo sacs are 8- nucleate and arise from one of the two dyad
cells formed after meiosis I.

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Tetrad formed as a result of meiosis in MMC, has four genetically different nuclei. Being a
derivative of two meiotic products, nuclei of a bisporic embryo sac are genetically
different (four nuclei are of one type and other four of a different type). A bisporic embryo
sac was first described in Allium fistulosum (Strasburger, 1879) and has since been
confirmed in several species of this genus.

Bisporic embryo sacs are also of two types.

1. Allium type
2. Endymion type
(i) Allium type (8 nucleate): In this type chalazal dyad cell participates in the
formation of the embryo sac (Figure 3.16).

Chalazal dyad

Fig. 3.16 Allium type embryo sac

(ii) Endymion type (8 nucleate): In this type micropylar dyad cell participates in the
formation of the embryo sac (Figure 3.17).

Micropylar dyad

Fig. 3.17-Endymion type embryo sac

3. Tetrasporic Type:

This type of embryo sac development is remarkable because of the complete elimination of
wall formation during meiosis and the participation of all four megaspore nuclei in the
formation of the embryo sac. That’s why it is called tetrasporic embryo sac. In this type
neither of the meiotic division (first as well as second) is accompanied by wall formation
so at the end of meiosis all the four haploid nuclei remain in a common cytoplasm forming
a coenomegaspore (four nuclei inside a cell) and all the four nuclei of the
coenomegaspore take part in the formation of embryo sac. This type of embryo sac is more

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heterogenous than a bisporic one because all the four nuclei of the coenomegaspore,
products of meiosis are genetically different. Three main types of patterns for embryo
sac development can become clear by going through Figure 3.18.

Figure 3.18-Three main types of embryo sac development: monosporic, bisporic, tetrasporic

Nuclear behaviour in tetrasporic embryo sac is variable. The arrangement of the four
nuclei in the coenomegaspore, before the beginning of post-meiotic mitosis, is of three
types.

1. 2+2 arrangement: two nuclei at the micropylar end and two nuclei at the chalazal
end (e.g. Adoxa –type)
micropylar end

chalazal end

2. 1+1+1+1 arrangement: one nucleus at the micropylar end, one at the chalazal
endand two placed laterally, one on each side. (e.g. Penaea type, Plumbago type,
Peperomia type)
micropylar end

chalazal end

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3. 1+3 arrangement: one nucleus at the micropylar end and three at the chalazal end
(e.g. Drusa type, Fritillaria type, Plumbagella type)
micropylar end

chalazal end

According to P. Maheshwari (1950), there are 7 types of tetrasporic embryo sacs-

1. Adoxa type
2. Penaea type
3. Plumbago type
4. Peperomia type
5. Drusa type
6. Fritillaria type
7. Plumbagella type
Depending on the following features tetrasporic embryo sacs are of many
types:

1. whether nuclear fusion occurs or not,

2. number of the post-meiotic mitoses in the coenomegaspore and
3. final organization of the embryo sac
I-No nuclear fusion occurs

1. Adoxa-type: 2+2 arrangement: two nuclei at the micropylar end and two nuclei at the
chalazal end. The embryo sac is eight nucleate, formed after single post-meiotic mitosis,
having similar organization to that of the Polygonum-type. Examples- In Adoxa,
Sambucus, Ulmus in the dicotyledons and to certain species of Erythronium, and Tulipa in
monocotyledons.

2. Penaea-type: 1+1+1+1 arrangement: one nucleus at the micropylar end, one at the
chalazal endand two placed laterally, one on each side. As a result of two post-meiotic
mitosis in the coenomegaspore, sixteen nucleate embryo sac is formed. The mature embryo
sac has 4 groups of 3 cells each, one group is at the micropylar end, one at the chalazal end
and two arranged laterally. The remaining four nuclei at the center are polar nuclei. The
micropylar triad functions as the egg apparatus.

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3. Plumbago-type: 1+1+1+1 arrangement: one nucleus at the micropylar end, one at

the chalazal end and two placed laterally, one on each side. In this type only one
post-meiotic mitosis occurs but the organization of mature embryo sac is different from
that in Adoxa-type. The mature embryo sac has an egg cell and a four- nucleate central cell.
The other three nuclei are cut-off as peripheral cells.

4. Peperomia type: 1+1+1+1 arrangement: one nucleus at the micropylar end, one at
the chalazal endand two placed laterally, one on each side. Embryo sac is 16 nucleate, like
Penaea-type (two post-meiotic mitosis) but here an egg apparatus comprising an egg and
only one synergid, six peripheral cells and a central cell with eight polar nuclei.

5. Drusa-type: 1+3 arrangement: one nucleus at the micropylar end and three at the
chalazal end. Embryo sac is also 16 nucleate (two post-meiotic mitosis) but the mature
embryo sac has a normal egg apparatus (3-celled), two polar nuclei, and 11 antipodal cells.

II-After the second meiotic division three chalazal megaspore nuclei fuse to
form a triploid nucleus of the coenomegaspore. The fourth nucleus at the
micropylar end remains haploid.

6. Fritillaria-type: 1+3 arrangement: one nucleus at the micropylar end and three at the
chalazal end.The true sequence of events in the embryo sac development of Lilium and
Fritillaria was fully understood on the basis of investigations of Bambacioni (1928) and
Cooper (1935).

In Lilium and Fritillaria the four megaspore nuclei behave in peculiar and distinctive
manner. Three of these nuclei migrate to the chalazal end of the sac and the remaining
nucleus is situated at the micropylar end.

Steps:

-The 1+3 arrangement represents the first, four-nucleate stage in the development of
embryo sac.

-The micropylar nucleus divide to form two haploid nuclei, the three chalazal nuclei fuse
(triploid) and subsequently divide to form two triploid nuclei. As a result, a second,
four-nucleate stage is observable, consisting of two haploid micropylar nuclei and two
triploid chalazal nuclei.

-A final (fourth) nuclear division occurs, producing four haploid micropylar nuclei (a
micropylar quartet) and four triploid chalazal nuclei (a chalazal quartet).

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Fig. 3.19-Types of embryo sacs in angiosperms (Modified from Maheshwari, Bot.Rev.14:1, 1948)

• The nuclei of the bisporic and tetrasporic embryo sacs are genetically not identical as they
are in monosporic embryo sacs, because they arise from two or four different meiotic
products.
II-Variation in the formation of meiotically unreduced (diploid) female
gametophytes during gametophytic apomixis (Drews and Koltunow 2011)

We know that there are two different forms of apomixis in angiosperms and these are:

1. Sporophytic apomixis

Here somatic cell of an ovule adjacent to a developing embryo sac develops directly into an
embryo. So we can say that sporophytic apomixis bypasses female gametophyte formation.
Example- in Citrus and mango.

2. Gametophytic apomixis

It involves formation of a meiotically unreduced (i.e. diploid) female gametophyte. The

diploid egg cell then forms an embryo by parthenogenesis (i.e. without fertilization) and
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