GINKGOALES

The order Ginkgoales is today represented by only one living member, i.e. Ginkgo biloba.
Ginkgoales was, however, very abundantly represented in the world by several species of
about 16 genera during the Triassic period of Mesozoic age, i.e. about 200,000,000 years ago.
Today, all the genera, except Ginkgo biloba, are extinct.
G. biloba is represented by five varieties viz. Ginkgo biloba var. aurea (Nelson) Beisson,
G.biloba var. fastigata Henry, G.biloba var. paciniata Carriere, G. biloba var. pendula Carnere
and G. biloba var. variegata Carriere. Due to the presence of a number of primitive
characters, as well as because of its long geological records, Ginkgo is called a “living
fossil”.
Details of the geological history of Ginkgoales indicate that its members started appearing on
the earth during Permian, achieved luxuriance and worldwide distribution during Triassic and
Jurassic periods of Mesozoic age, started fading out of existence during Cretaceous and now
represented only in some parts of Southern and Eastern China by only one living member i.e.
Ginkgo biloba.
Ramanujam (1976), while tracing the geological history of Ginkgoales, reported that a few
records of this order are known from Late Palaeozoic of India. These include Ginkgophyton,
Psygmophyllum and Rhipidopsis.
Seward (1938) considered Ginkgo “as one of the wonders of the world” and stated that has
persisted with little change until the present through a long succession of ages when the earth
was inhabited by animals and plants”. Arnold (1947) mentioned that “Ginkgo biloba is one of
the oldest living plants and may indeed be the oldest living genus of the seed plants”
Distribution of Ginkgo:
Ginkgo biloba is a tall slender and beautiful tree. It is commonly called Maiden-hair Tree
because its new leaves resemble very much like those of Adiantum (called maiden hair fern)
both in form and venation. Ginkgo still exists in the wild state in South-eastern China, along
“the north western border of Chekiang and south eastern Anhwei”. In China and Japan it is
grown as a sacred tree in temple gardens. It is cultivated in the United States as a shade tree.
It is also successfully cultivated in some gardens of Europe, America and India.
1. Tall, well-branched trees with short and long shoots. However, some earliest fossil
members were without short and long shoots.
2. Wood is pycnoxylic.
3. Leaves are large, leathery and fan-shaped or strap-shaped. They are often deeply divided.
4. Dichotomous venation is usually present in the leaves.
5. Un-branched, catkin-like male organs are axillary in position.
6. Male organs bear micro-sporangiophores.
7. Each micro-sporangiophore possesses 2-12 pendulous microsporangia.
8. Spermatozoids are motile and contain spiral bands of flagella.
9. Ovules are terminal in position on branched or un-branched axillary axes. They are 2-10 in
number.
10. Seeds are large-sized.
11. Each seed contains a fleshy outer layer and a middle stony layer.
Morphological Features of Ginkgo:
The plant body of Ginkgo biloba is sporophytic, and the sporophyte resembles several
conifers in general habit. The trees have a pronounced ex-current habit of growth and attain a
height up to 30 metres. A very irregular pattern of branching is shown by Ginkgo trees.
The branches are dimorphic i.e. bear long shoots which are of unlimited growth with
scattered leaves and dwarf shoots which are short branches of limited growth.
Long shoots elongate rapidly, sometimes as much as 50 cm in a year. Dwarf shoots grow
rather slowly. A dwarf shoot of 2-3 cm length may be several years old. Sometimes there is
no clear-cut distinction between two types of shoots, and the dwarf shoot may convert into a
long shoot while the latter may sometimes convert into a dwarf shoot for a year or two and
then revert back into a long shoot.
The foliage leaves, present on the long shoots, are deeply lobed while those on the dwarf
shoots are not so deeply lobed and sometimes more or less entire (Figs. 10.4, 10.5). Bierhorst
(1971) opined that differences in the growth pattern of long and dwarf shoots may be due to
the quantities of auxin produced in the apical meristems.

Ginkgo biloba possesses a long tap root system. The roots are extensively branched and
penetrate deep into the soil. The foliage leaves are simple,large, petiolate and wedge-shaped
or fan-shaped with expanded apex and narrow base.
They resemble Maiden-hair fern (Adiantum) and by this they can be distinguished among
gymnosperms In general, the leaves are bilobed, and hence the name ‘biloba’ was suggested
to the species by Linnaeus (1771).
However, the leaves with many lobes (Fig 10.6) are also present invariably on the tree.The
leaves of the long shoot and seedlings are deeply bilobed while that of the dwarf shoot are not
so deeply lobed and may be sinuate or even entire. They may be pale yellow, golden yellow
or dark green in colour. A typical dichotomous type of venation (Fig. 10.7) is present in the
leaves.
Foliar epidermis also exhibits some distinguishable characters in Ginkgo biloba. The
epidermal cells are rectangular over the veins while polygonal in outline in between the
veins. The leaves are hypostomatic (i.e. bear stomata only on the lower surface of the leaf).
Kanis and Karstens (1963), however, observed a few stomata on the upper surface of the
leaves on long shoot, thus making them amphistomatic. Stomata remain surrounded by 4-6
subsidiary cells with finger-like outgrowths overarching the guard cells.
5. Anatomy of Ginkgo:
(i) Root:
In transverse section (Fig. 10.8) the roots are somewhat circular in outline. Mature roots are
surrounded by phellogen or suberized cells of cortex. A large portion of the young root is
occupied by multilayered, thin-walled cortex which contains several tannin- filled cells and
calcium oxalate crystals. Mucilage canals are also prominently visible (Fig. 10.8)

In young roots, a layer of endodermis and uni-layered pericycle are clear. Mature
roots,however, lack such a distinction. Diarch condition is clearly visible in the young roots.
Xylem is exarch. It remains separated by the phloem strands. Sometimes the roots also show
triarch condition.
(ii) Stem:
The young stem (long shoot) is more or less circular in outline and remains surrounded by a
single-layered, thickly circularized epidermis (Fig. 10.9) made of brick-shaped cells.
Epidermis is replaced by periderm in the older stems.

Inner to the epidermal layer is present a well-marked region of parenchymatous cortex. It

contains mucilaginous canals, sphaeraphides and many tannin-filled cells. Cortex is
comparatively less extensive in long shoots than dwarf shoots. Endodermis and pericycle are
not well-marked in long shoot.
Several conjoint, collateral, open and endarch vascular bundles are arranged in a ring in very
young stem. Two leaf traces, one for each leaf, are given out. After the onset of the secondary
growth, the vascular cylinder of the stem becomes an endarch siphonostele with no
parenchyma in the wood except that of uniseriate medullary rays.
Protoxylem has spiral thickenings while bordered pits are present on the radial walls of the
metaxylem tracheids. Sieve tubes and phloem parenchyma constitute the phloem. A narrow
pith, containing mucilage canals and sphaeraphides, is present in the centre of long shoot,
while in dwarf shoot the pith is comparatively more extensive.
Secondary Growth in Stem:
Cork cambium cuts periderm towards outer side and secondary cortex towards inner side
(Fig. 10.10) Periderm replaces the epidermis. Mucilage canals are absent.

A single ring of cambium remains active throughout and cuts secondary phloem towards
outer side and secondary xylem towards inner side. Crushed patches of primary phloem
towards outer side and primary xylem towards inner side are present. Secondary phloem
consists of sieve tubes and phloem parenchyma. Secondary xylem consists of tracheids. Ill-
defined annual rings are also seen.
Study of the tangential longitudinal sections of long and dwarf shoots shows that uniseriate
medullary’ rays are 1 -3 cells in height in long shoot (Fig. 10.11) while 1-15 cells in height in
dwarf shoot (Fig. 10.12).
One or two rows of bordered pits are present on the radial walls of the tracheids (Fig. 10.13).
Pits are circular in outline and have a clear torus. Bars of Sanio are also present. Bars of
Sanio do not occur in primary wood. Trabeculae of Sanio, which cross the lumen of the
tracheids, are also present.

(iii) Leaf:
A layer of epidermis is present on upper as well as lower sides of leaf. The epidermis is
thickly cuticularised and consists of rectangular to polygonal cells. Haplocheilic type of
stomata, restricted only to the lower epidermis, are present. Kanis and Karstens (1963),
however, reported some stomata on the upper epidermis of the leaves on long shoots of male
plants only.
Mesophyll is present in between the two epidermal layers. It is not well-differentiated into
palisade and spongy parenchyma (Fig. 10.14). The leaves of long shoot, however, show a
distinct palisade region. Many mucilage canals or secretory canals and a few tannin-filled
cells are also present in the mesophyll region.
Vascular bundles of the vein have an indistinct mesarch structure (Fig. 10.14). Each vascular
bundle may be surrounded by a sclerenchymatous bundle sheath in mature leaves.
(iv) Petiole:
The petiole remains covered by a layer of thickly cuticulanzed epidermis, the continuity of
which is broken by stomata. Inner to the epidermis are present a few hypodermal layers. Few
mucilage canals, tannin-filled cells and sphaeraphides are irregularly distributed in the cortex.
The petiole is supplied by a pair of endarch vascular bundles (Fig. 10.15).
The detailed structure of a vascular bundle is shown in Fig. 10.16. It remains surrounded by a
sclerenchymatous bundle sheath. Protoxylem has spiral thickenings. Cambium is clearly
visible Uniseriate rays are present in the xylem. These are continuous with those of the
phloem.

6. Reproductive Structures of Ginkgo:

Ginkgo biloba is dioecious. Male and female plants are,however, difficult to be differentiated,
when young. According to Lee (1954) sex in Ginkgo is determined by sex chromosomes (XY
in male and XX in female). Reproductive bodies of Ginkgo are most primitive among living
seed plants except some Cycadales.
(i) Male Strobilus:
Male or microsporangiate fructifications develop in catkin-like clusters (Fig. 10.17 A) on the
dwarf shoots of male frees.Each male strobilus contains several microsporophyll’s arranged
loosely on a central axis (Fig. 10.17B). Each microsporophyll has a long stalk terminating
into a hump or knob.
It contains two pendant microsporangia (Fig. 10.17C). According to some workers this
terminal knob represents an abortive sporangium. A mucilage duct is present in the knob.
Rarely more than two sporangia are present in a microsporophyll.
Each sporangium is a tubular structure surrounded by many layers (Fig. 10.17D). The
outermost layer of sporangial tissue differentiates into a tapetum. The sporogenous cells of
the sporangium undergo reduction division and form many haploid microspores Wolniak
(1976) has studied the ultrastructure of the microspore mother cell.
(ii) Development of Microsporangium:
It is of eusporangiate type, i.e., single archesporial cell divides by a periclinal wall forming
primary wall cell and primary’ sporogenous cell (Fig. 10.18A,B). The former develops into
wall of microsporangium while the latter develops into sporogenous tissue. Sporangium
dehisces by means of a longitudinal slit.
(iii) Female Strobilus:
The female strobili develop in groups in the axil of leaves or scaly leaves present on the
dwarf shoots (Fig. 10.19). These are much reduced structures, each having a long stalk or
peduncle, which bifurcates apically. Each bifurcation usually bears a single ovule. Out of the
two sessile ovules one generally aborts earlier.
Rarely, three, four or more ovules on a peduncle are also observed. Sprecher (1907) reported
an axis with seven ovules, each borne singly on its independent peduncle, and not in pairs. At
the base of each ovule is present a fleshy cup-like structure called ring or collar. The leaves
surrounding the ovules do not show their bilobed character.

The development of ovule, mega-sporogenesis and structure of the mature ovule is similar to
that of Cycas. There is a thick integument consisting of three layers, i.e., outer fleshy, middle
stony and inner fleshy layers (Fig. 10.20). A major difference between Ginkgo and Cycas is
that the integuments of Ginkgo do not have any noticeable vascular supply.
Each ovule has a large and prominent nucellus. Free apex of the nucellus breaks down into a
pollen chamber (Fig. 10.21) forming the nucellar beak. Quite deep in the nucellus tissue, a
functional spore mother cell becomes prominent. The spore mother cell develops into a
tetrad, of which only the innermost megaspore remains functional and develops into female
gametophyte.
The vascular supply of the ovule is ill-developed. Two vascular strands enter the inner fleshy
layer and reach up to the free part of the nucellus without branching. Outer fleshy layer lacks
any prominent vascular supply.
7. Gametophytes in Ginkgo:
(i) Male Gametophyte:
Major aspects of the development of male gametophyte of Ginkgo biloba (Fig 10.22) are
similar to that of Pinus. Male gametophyte starts developing in situ, i.e. within the
sporangium.
Friedman (1987) studied the growth and development of the male gametophyte of Ginkgo
biloba using three-dimensional computer reconstruction (Fig. 10.22). Pollen germination
starts with a diffuse swelling of tube cell. Tube cell becomes very elaborate with a proximal
un-branched tube and a much branched haustorial system.

Microspore is the first cell of the male gametophyte. Each microspore is a rounded structure
having thin intine and thick exine layers. A centrally located nucleus contains one or two
nucleoli and remains surrounded by dense cytoplasm. An un-thickened portion, called pore,
is also present in each microspore. It is a region where exine is not covering the intine.
Two unequal cells are formed by the first mitotic division in the microspore. These are called
1st prothallial cell and inner cell (Fig. 10.23 A). The 1st prothallial cell is smaller than that of
inner cell. The inner cell again divides forming a 2nd prothallial cell and an antheridial initial
(Fig. 10.23B).
By this time the 1st prothallial cell starts to degenerate. The antheridial initial then divides
forming a generative cell near the 2nd prothallial cell and a tube cell (Fig. 10.23D). The tube
cell does not divide any further. This is the 4-celled stage of the male gametophyte and the
microspores are dispersed by wind at this stage.

Pollination in Ginkgo biloba takes place sometime in April. At the time of pollination the
ovule secretes a mucilaginous substance. Further development of the male gametophyte takes
place in the pollen chamber. In pollen chamber the generative cell of the 4-celled young male
gametophyte divides into a stalk cell, which is close to the 2nd prothallial cell, and a body
cell.
Now the intine protrudes out near the pore and functions as a pollen tube (Fig. 10.23E). The
pollen rube swells up and advances towards the archegonia by protruding into the nucellus.
Just before fertilization the body cells divides into two, and the wall formation is at right
angle to the base of pollen grains.
The blepharoplasts appear on the body of these so-formed two cells which function as sperms
or male gametes (Fig. 10.23F). The structure of these multi-flagellated sperms, as well as
their behaviour in the tube at the time of fertilization has been investigated by
Shimamura(1937), Lee (1955) and Favre-Duchartre (1956).
Motile sperms in gymnosperms were observed for the first time on September 9, 1896 by
Hirase, and his this discovery is considered as one of the most remarkable events in plant
morphology. The ultrastructure of male gamete of Ginkgo has been studied by Gifford and
Lin (1975).
(ii) Female Gametophyte:
The development of the ovule and female gametophyte in Ginkgo biloba has been studied by
Carothers (1907). Out of the four megaspores formed from the megaspore mother cell, only
the lowermost remains functional and the remaining three degenerate (Fig. 10.24) A).
The nucleus of the functional megaspore migrates towards the micropylar end and divides
into two followed by a number of free-nuclear divisions forming hundreds of free nuclei.
A large vacuole is present in the megaspore at this stage (Fig 10.24B). Now the wall
formation starts (Fig. 10.24B,C) and progresses from the periphery towards the centre. Dunng
wall formation first the anticlinal walls are formed followed by the vertical walls.
Each cell generally contains one nucleus but in some cells 2-3 nuclei are also seen. Due to
more rapid cell divisions on the micropylar end a pole-like structure develops. This is called
tent-pole (Fig. 10.24D). The female gametophyte possesses abundant chlorophyll.
The development of archegonium starts from the cells towards the micropylar end of the
female gametophyte. According to Favre-Duchartre (1958) the archegonial initials start
appearing sometime in June.
Each archegonium possesses a short neck made up of only four cells and a small venter
having a central cell. The central cell later on forms a ventral canal cell and an egg cell (Fig.
10.25). According to Bierhorst (1971) only two cells are present in the archegonial neck.

Ultra structural study of the female gametophyte of Ginkgo biloba has been made by
Dexheimer (1973). He observed four zones based on food reserves.
These are:
(1) Lipid zone,
(2) Starch proteolipid zone with large vacuolated cells,
(3) Starch zone made of large vacuolated cells with peripheral cytoplasm, and
(4) Deep or central zone with very little reserve contents.
Fertilization:
The pollen tube reaches up to the neck of the archegonium just after its (archegonium)
differentiation. The tube ruptures releasing the sperms and the other contents in the
archegonial chamber. A sperm passes through the neck of the archegonium, comes in contact
and fuses with the egg nucleus exactly in the same way as in Cycas.
8. Embryogeny in Ginkgo:
The zygote becomes enlarged and starts dividing by many free-nuclear divisions just after
fertilization. The nuclei are distributed irregularly (Fig. 10.26A). The cell wall formation
starts just after a few divisions of the nucleus and soon a pro-embryo is differentiated (Fig.
10.26B). Three types of cells are clearly differentiated in the pro-embryo (Fig. 10.26C).
The cells towards the chalazal end are smaller in size with dense cytoplasm, followed by
another zone of comparatively larger cells, which do not possess such a dense cytoplasm.
Elongated cells are present in the uppermost zone. There is no differentiation of suspensor in
the embryo.
The development of the cotyledon starts from the lowermost zone soon after the
differentiation of these zones (Fig 10.26D,E). Roots, stem and cotyledons are soon
differentiated (Fig. 10.26 E,F). Generally, five leaves are present in the mature embryo. Out
of these five leaves the first two are decussate with cotyledons while the remaining leaves are
irregular in arrangement in embryo and seedling.

The germination of the seed is of hypogeal type and quite similar to that of Cycas. A strong
tap root develops soon and the seedling bears many bilobed leaves, which is characteristic
feature of Ginkgo biloba. Soon, a mature plant, with many more leaves, develops.
In a recent study, Dogra (1992) opined that development of embryo in Ginkgo biloba is a
continuous process from the time of fertilization until the seed germination. Actually, the
major part of the growth of embryo takes place when the seed is detached from the tree and is
lying on the ground.
9. Economic Importance of Ginkgo:
Ginkgo biloba tree is treated as a sacred plant in China and Japan and worshipped by the
people. It is regarded as “Holy Tree” by Buddhist monks and nuns because of its tall, strong
and majestic appearance. It is grown as an ornamental plant in gardens in several countries.
In autumn, the green leaves turn yellow, showing a magnificent autumn scenery. The
endosperm of its roasted seeds is edible, but seeds may prove fatal if taken in large quantity.
After the outer skin of its apricot-like seeds is peeled off or decayed, the kernel left is called ”
White Fruit”. Ginkgo is, therefore, also named “White fruit tree”.
The kernel is also used as food and Chinese medicine as well. If it is taken raw, it functions
as detoxicating and asthma-curing medicine. Although the plants are highly resistant to
pathogenic infections, specially of bacteria and fungi, their wood is not of much commercial
importance. It also possesses a strong capacity of resistance to air pollution by smoke and
poisonous gases.
Since an unpleasant odour is emitted from the ripe seeds of female trees, male trees are
largely preferred and grown as shade trees in China, Japan and United States. Ginkgo trees
grow and develop very slowly.
It is said that the tree planted by grandfather will bear “fruit” at the time when his grandson
grows up. Therefore, it is also known as “grandfather grandson tree”.An extract of leaves of
G. biloba is useful in the treatment of cerebral insufficiency and vertigo.

Phylogeny of Ginkgoales:
Some of the striking peculianties of Ginkgo biloba, the only living representative of
Ginkgoales, include:
(i) Bilobed, fan-shaped leaves with dichotomous open venation,
(ii) Ovule bearing a collar at the base,
(iii) Micro-sporangiophore bearing a hump-like outgrowth at its apex,
(iv) Presence of tent pole at the tip of the female gametophyte, and
(v) Absence of suspensor in its embryo.
In-spite of these characteristics, Ginkgoales resemble in some or other aspects with
Cordaitales, Pteridospermales, Filicales, Cycadales and even Coniferales.

Resemblances with Cordaitales

(i) Presence of double leaf trace,
(ii) Endospermic beak in the mature ovule, and
(iii) The probable motility of the spermatozoids of the members of both Ginkgoales and
Cordaitales are some of the resemblances between these two groups.
Resemblances with Pteridospermales:
(i) Presence of leaf gap in the stem,
(ii) Dichotomous open venation in the wedge-shaped leaves of Ginkgo and pinnules of some
seed ferns (Ptendospermales),
(iii) A distinct pollen chamber,
(iv) A massive fleshy layer in the ovules, and
(v) Collar at the base of the ovules in Ginkgo and cupule in ovules of some Pteridospermales,
are some of the characteristics showing resemblance between Ginkgoales and
Pteridospermales.
Resemblances with Filicales:
(i) The similarities between the primary xylem structure, secondary xylem structure and
periderm of Ginkgo and some ferns (Ophioglossum and Botrychium),
(ii) Dichotomous open venation in their leaves,
(iii) Multi-flagellated and motile spermatozoids, and
(iv) Presence of ventral canal cell in the archegonia in some of their members.
Resemblances with Cycadales:
(i) Presence of multi-flagellated spermatozoids,
(ii) Well-developed nucellar beak and pollen chamber,
(iii) Haustorial nature of pollen tube,
(iv) Presence of large egg, massive female gametophyte and well-developed venter,
(v) Endoscopic embryo with two cotyledons,
(vi) Seed with thick and well-developed integument, and
(vii) Hypogeal type of seed germination.
In-spite of these above-mentioned similarities, there are several major differences between
Ginkgoales and Cycadales which put them quite apart from each other.
They differ from each other in the:
(i) General structure of their male and female reproductive organs,
(ii) Vascular supply of their ovules,
(iii) Development of their male gametophytes,
(iv) Branched (Ginkgoales) and generally un-branched (Cycadales) nature of their stems,
(v) Simple (Ginkgoales) and compound (Cycadales) nature of their leaves, and
(vi) Presence (Cycadales) or absence (Ginkgoales) of circinate venation in their leaves, etc.
Resemblances with Coniferales:
Ginkgo has several characteristics in common with many members of Coniferales which
compel one to think Ginkgoales as well-defined Coniferophytes. Florin opined that all
Ginkgoales, Coniferales, Cordaitales and Taxales belong to the same natural group named as
Coniferopsida.
All these four orders of Coniferopsida constitute parallel lines of evolution and separated
from each other in the Upper Devonian or Carboniferous times according to Florin.
Some major characteristics common to both Ginkgoales and Coniferales
are listed below:
(i) General appearance of tree is cone like,
(ii) Extensively branched stem with two types of branches, i.e. long shoots and dwarf shoots,
(iii) Leaves are simple with sunken stomata,
(iv) Cortex and pith are narrow and wood is pycnoxylic,