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SEXUAL REPRODUCTION IN FLOWERING PLANTS

Pre-Fertilisation: Structures and Events


Much before the actual flower is seen on a plant, the decision that the plant is going to flower has taken place. Several hormonal and structural changes are initiated which lead to the differentiation and further development of the floral primordium. Inflorescences are formed which bear the floral buds and then the flowers. In the flower the male and female reproductive structures, the androecium and the gynoecium differentiate and develop.
You would recollect that the androecium consists of a whorl of stamens representing the male reproductive organ and the gynoecium represents the female reproductive organ.

Stamen, Microsporangium and Pollen Grain

Figure 2a shows the two parts of a typical stamen – the long and slender stalk called the filament, and the terminal generally bilobed structure called the anther. The proximal end of the filament is attached to the thalamus or the petal of the flower. The number and length of stamens are variable in flowers of different species. If you were to collect a stamen each from ten flowers (each from different species) and arrange them on a slide, you would be able to appreciate the large variation in size seen in nature. Careful observation of each stamen under a dissecting microscope and making neat diagrams would elucidate the range in shape and attachment of anthers in different flowers.
A typical angiosperm anther is bilobed with each lobe having two theca, i.e., they are dithecous (Figure 2). Often a longitudinal groove runs lengthwise separating the theca. Let us understand the various types of tissues and their organization in the transverse section of an anther (Figure 3 a). The bilobed nature of an anther is very distinct in the transverse section of the anther. The anther is a four-sided (tetragonal) structure consisting of four microsporangia located at the corners, two in each lobe.


(a) A typical stamen; (b) three–dimensional cut section of an anther

(a) Transverse section of a mature anther; (b) Enlarged view of one microsporangium showing wall layers; (c) A dehisced anther

The microsporangia develop further and become pollen sacs. They extend longitudinally all through the length of an anther and are packed with pollen grains.
Structure of microsporangium: In a transverse section, a typical microsporangium appears near circular in outline. It is generally surrounded by four wall layers (Figure 3 b)– the epidermis, endothecium, middle layers and the tapetum.
The outer three wall layers perform the function of protection and help in dehiscence of anther to release the pollen. The innermost wall layer is the tapetum. It nourishes the developing pollen grains. Cells of the tapetum possess dense cytoplasm and generally have more than one nucleus. Can you think of how tapetal cells could become bi-nucleate?
When the anther is young, a group of compactly arranged homogenous cells called the sporogenous tissue occupies the centre of each microsporangium.
Microsporogenesis : As the anther develops, the cells of the sporogenous tissue undergo meiotic divisions to form microspore tetrads. What would be the ploidy of the cells of the tetrad?
As each cell of the sporogenous tissue is capable of giving rise to a microspore tetrad. Each one is a potential pollen or microspore mother cell (PMC). The process of formation of microspores from a pollen mother cell through meiosis is called microsporogenesis. The microspores, as they are formed, are arranged in a cluster of four cells–the microspore tetrad (Figure 3 a). As the anthers mature and dehydrate, the microspores dissociate from each other and develop into pollen grains (Figure 3 b). Inside each microsporangium several thousands of microspores or pollen grains are formed that are released with the dehiscence of anther (Figure 3 c).
Pollen grain: The pollen grains represent the male gametophytes. If you touch the opened anthers of Hibiscus or any other flower you would find deposition of yellowish powdery pollen grains on your fingers. Sprinkle these grains on a drop of water taken on a glass slide and observe under a microscope. You will really be amazed at the variety of architecture –sizes, shapes, colours, designs – seen on the pollen grains from different species (Figure 4).

Scanning electron micrographs of a few pollen grains

Pollen grains are generally spherical measuring about 25-50 micrometers in diameter. It has a prominent two-layered wall. The hard outer layer called the exine is made up of sporopollenin which is one of the most resistant organic material known. It can withstand high temperatures and strong acids and alkali. No enzyme that degrades sporopollenin is so far known. Pollen grain exine has prominent apertures called germ pores where sporopollenin is absent. Pollen grains are wellpreserved as fossils because of the presence of sporopollenin. The exine exhibits a fascinating array of patterns and designs. Why do you think the exine should be hard? What is the function of germ pore? The inner wall of the pollen grain is called the intine. It is a thin and continuous layer made up of cellulose and pectin. The cytoplasm of pollen grain is surrounded by a plasma membrane. When the pollen grain is mature it contains two cells, the vegetative cell and generative cell (Figure 5b). The vegetative cell is bigger, has abundant food reserve and a large irregularly shaped nucleus. The generative cell is small and floats in the cytoplasm of the vegetative cell. It is spindle shaped with dense cytoplasm and a nucleus. In over 60 per cent of angiosperms, pollen grains are shed at this 2-celled stage. In the remaining species, the generative cell divides mitotically to give rise to the two male gametes before pollen grains are shed (3-celled stage).


(a) Enlarged view of a pollen grain tetrad; (b) stages of a microspore maturing into a pollen grain

Pollen grains of many species cause severe allergies and bronchial afflictions in some people often leading to chronic respiratory disorders– asthma, bronchitis, etc. It may be mentioned that Parthenium or carrot grass that came into India as a contaminant with imported wheat, has become ubiquitous in occurrence and causes pollen allergy.
Pollen grains are rich in nutrients. It has become a fashion in recent years to use pollen tablets as food supplements. In western countries, a large number of pollen products in the form of tablets and syrups are available in the market. Pollen consumption has been claimed to increase the performance of athletes and race horses (Figure 6).

Pollen products

When once they are shed, pollen grains have to land on the stigma before they lose viability if they have to bring about fertilisation. How long do you think the pollen grains retain viability? The period for which pollen grains remain viable is highly variable and to some extent depends on the prevailing temperature and humidity. In some cereals such as rice and wheat, pollen grains lose viability within 30 minutes of their release, and in some members of Rosaceae, Leguminoseae and Solanaceae, they maintain viability for months. You may have heard of storing semen/ sperms of many animals including humans for artificial insemination. It is possible to store pollen grains of a large number of species for years in liquid nitrogen (-196°C). Such stored pollen can be used as pollen banks, similar to seed banks, in crop breeding programmes.

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