CLASS HEPATICAE ORDER MARCHANTIA: the best known species of thalloid liverworts are in the genus Marchantia.

The thalles are 30 cells thick in the center and 10 cells thick at the margin and forks dichotomously as it grows. Each branch has a notch at the apex and central groove that extends back lengthwise behind the notch. The thalli grow as meristematic cells as the notches divide. Older tissues at the rear decay as new growth is added. The upper surface of the thalles is divided into diamond-shaped or polygonal segment, with the segments line, marking the limits of the opening into the interior. Through a microscope, a sectional liverworts looks like a series of covered cactus garden sitting on a decorative rock wall. The rock wall may comprise mostly of thalles consisting of parenchyma cells that have few (if any) chloroplasts. The tissue apparently stores substances produced in other cells. The bottom layer of the cells is an epidermis from which rhizoids and scales arise. The cactus garden consists of upright branching rows of chlorenchyma cells on the air space. Vertical walls enclose the individual gardens which are covered by a slightly dome-shaped layer of epidermal cells. The conspicuous pore which remains open at some times is located in the center of each roof and looks like a short suspended open barrel. The pore is like the stomata of higher plants i.e. allows for aeration of the thalles with a minimum dehydration but it is incapable of significant appearance in its aperture. ASEXUAL REPRODUCTION: Marchantia reproduces asexually by gemmae (singular gemma). Gemmae are tiny lens-shaped pieces of tissue that become detached from the thalles. They are produced in gemmae cups scattered over the upper surface of the liverwort gametophyte. Raindrops may splash the gemmae ass much as 1m away. While the gemmae are in the cup, lunaric acid (an endogenous growth regulator found in the liverworts) inhibits their further development until they come out i.e. each is capable of growth into a new thalles as soon as they come out. In addition, parts of the older thalles may die isolating patches of active tissue which may then continue to grow independently. SEXUAL REPRODUCTION: Marchantia is dioecious i.e. the male and female plants are distinct and separate. The gametangia which are produced on separate male and female plants gametophytes are more specialized than those of other liverworts. Both types of gametangia are formed on gametangiophores which are umbrella-like structures borne on slender stalks arising from the central grooves of the thalles. The top of the male gametangiophores termed antheridiophore which is disc-like with a scalloped margin while that of the female gametangiophore is termed archegoniophore which also looks like a hub and spokes of a wagon wheel (like bicycle). Antheridia which are club-shaped make gametangia containing numerous sperms and are produced in rows just beneath the upper surface of the antheridiophore.

Archegonia are flask-like female gametangia with each containing a single egg and are also produced in rows and hang neck downwards beneath the spokes of the archegoniophores. The sperms are extruded in a mucilaginous mass. Rain drops sometimes splash the released sperms which have flagella more than a meter away. Fertilization may occur before the stalks of the archegoniophore have finished growing. After fertilization, the zygote develops into a multicellular embryo (an immature sporophyte). A knob-like foot anchors the sporophyte (the diploid spore producing phase) in the tissues of the archegoniophore, the sporophyte hangs suspended by a short thick stalk called seta. The seta serves to lower the sporangium away from the archegonial disc and thus facilitate distribution of the spore. The main part of the sporophyte in which different types of tissue develop is called sporangium. Liverwort sporophytes typically have no stomata. Spore mother cells in the capsule undergo meiosis producing haploid spores. Other capsules cells do not undergo meiosis but remain diploid and develop rather into pointed elators which have spiral thicken and are sensitive to changes in humidity. Spore dispersal in Marchantia as the elaters twist and untwist rapidly. Until it is matured, the sporophyte is protected by a cap-like tissue i.e. the calyptra which grows out from the gametophyte and by other membranes covering the capsule. The capsule splits at maturity and air currents carry the spores away. Under favorable conditions, the spores germinate producing new gametophyte.

ORDER JUNGERIMA: This is the largest order of the liverworts containing about 2/3 of all species in the Hepaticae. They are often abundant in tropical jungles and fog belts. VEGETATIVE STRUCTURE: The gametophytes of these liverworts are leafy in contrast to the thalloid plant i.e. the gametophyte has stem-like and leaf-like organs that may be held somewhat erect off the ground. A very common and widespread genus is porella. Other examples are Funaria, Lophocoloa and Marsypella. Leafy liverworts always have two rows of partially overlapping leaves whose cells contain distinctive oil bodies. The leaves consist of a single layer of cells and have folds and lobes. In the tropics, the lobes form little water pockets in which tiny animals are always present. It has been suggested that these water pockets may function like the pitchers of the pitcher plant. A third row of under leaves called amphigastria which are smaller than the other leaves and not visible from the top is often present on the underside of leafy liverworts. Classification of the leafy liverworts is based largely on the arrangement and lobbing of the leaves. When the anterior margins of the leaves lie regularly beneath the posterior of the front,

the arrangement is said to be subcucous and when the convex (opposite) occurs, the arrangement is said to be incubous. A few rhizoids which anchor the plant develop from the stem-like axis and the base of the nearer leaves. SEXUAL REPRODUCTION: reproduction is essentially similar to that described for the Marchantia. Specialized gametangiophore are however never produced. Both monoecious and dioecious forms occur sometimes in the same genus. Antheridia usually occur singly in the axis of leaves of special branches of limited group. Archegonia are always restricted to the apices of stem and its branches. Because they are apical, the sporophyte terminates the growth of the main shoot so that vegetative growth is continued by lateral sympodial branching. Archegonia are generally enclosed within a cylindrical unistratos chlorophyllose tube, i.e. the perianth. The sporophytes resemble those of Marchantia. At maturity, the sporophyte capsule may be pushed out from among the leaves as the seta elongates. The capsule or the sporangium usually opens by 4 longitudinal lines. When the spore germinates, it produces the protonema which consists of a short filament of photosynthetic cells. The protonema soon develops into a new gametophyte.

ORDER PORELLA: Porella is the commonest acrogenous leafy liverwort. It grows on moist rocks, tree trunks and old walls, forming a compact greenish patch practically covering the medium on which it grows. STRUCTURE: the plant body consists of a slender dorsoventrally prostrate stem and leafy branches. The lower side of the stem bears several rhizoids primarily for anchorage. The leaves are arranged in three rows; 2 rows of dorsal leaves and a row of ventral small leaves called amphigastria. The dorsal leaves are unequally bilobed and overlapping. The plant grows by means of an apical tetrahedral cell which cuts off segments on 3 sides. REPRODUCTION: vegetative reproduction may take place by breaking off of some of the branches or by the formation of unicellular or multicellular gemmae on the margin or at the apex of the leaf. SEXUAL REPRODUCTION: porella is dioecious; the male plants are usually smaller and produce special short and lateral antheridial branches. These bear antheridia each in the axle of the leaves or bracts. Paraphyses (hair-like struct) may be present. The female plants are larger and produce larger archegonial branches. The archegonia on this are always borne terminally either singly or in a group. Paraphyses may be present. Each antheridium is globular barley surrounded by a wall or a jacket and provided with a long multicellular stalk. It is packed with antherizoid mother cells (androcytes), each giving rise to a minute bi-ciliate antherizoid. Each archegonium has a short multicellular stalk, a venter with an egg cell and egg nucleus; a ventral

canal cell, a long neck with 6 to 8 neck canal cells and a wall. The neck is nearly as broad as the venter. Fertilization takes place in the usual way. The antherizoids when liberated swim in water to the archegonium. They enter the apical opening and finally one of them fuses with the egg nucleus. The fertilized egg forms a zygote.

SPOROPHYTE: The zygote secretes a wall around itself and soon grows in size. It divides and redivides and soon gives rise to the sporophyte. This consists of a foot, seta and a capsule. The capsule is globuluse and surrounded by a wall (jacket) two or four layers thick. It encloses short spender spirally thickened elaters and many spores.. the sporophyte is surrounded by a calyptra, perianth and involucre. The calyptra is the envelope developed from the venter. The other two envelopes are formed by a united leaves or bracts. When matured, the capsule dehisces by 4 valves the spores are liberated. GERMINATION OF THE SPORES: Under favorable conditions, the spores germinate and give rise to a small multicellular body, the protonema. Soon, its apical cells become active and produces the shoot and leaves of anew porella plant.