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Learning Goals
By the end of this reading you should be able to:
- Explain why angiosperms are the dominant form of plant life in most terrestrial ecosystems
- Describe the main parts of a flower and their purposes
- Detail the life cycle of an angiosperm
- Discuss the two main groups of flowering plants
Introduction
Undisputed fossil records place the massive appearance and diversification of angiosperms in the middle to the late Mesozoic era. Angiosperms (“seed in a vessel”) produce a flower containing male and/or female reproductive structures. Fossil evidence indicates that flowering plants first appeared in the Lower Cretaceous, about 125 million years ago, and were rapidly diversifying by the Middle Cretaceous, about 100 million years ago. Earlier traces of angiosperms are scarce. Fossilized pollen recovered from Jurassic geological material has been attributed to angiosperms. A few early Cretaceous rocks show clear imprints of leaves resembling angiosperm leaves. By the mid-Cretaceous, a staggering number of diverse flowering plants crowd the fossil record. The same geological period is also marked by the appearance of many modern groups of insects, including pollinating insects that played a key role in ecology and the evolution of flowering plants (Fig. 1).
Although several hypotheses have been offered to explain this sudden profusion and variety of flowering plants, none have garnered the consensus of paleobotanists (scientists who study ancient plants). New data in comparative genomics have, however, shed some light on the evolution of angiosperms. Rather than being derived from gymnosperms, angiosperms form a sister clade (a species and its descendants) that developed in parallel with the gymnosperms.
From their humble and still obscure beginning during the early Jurassic period, the angiosperms—or flowering plants—have evolved to dominate most terrestrial ecosystems. With more than 250,000 species, the angiosperm phylum (Anthophyta) is second only to insects in terms of diversification. The success of angiosperms is due to two novel reproductive structures: flowers and fruit. The function of the flower is to ensure pollination. Flowers also provide protection for the ovule and developing embryo inside a receptacle. The function of the fruit is seed dispersal. They also protect the developing seed. Different fruit structures or tissues on fruit—such as sweet flesh, wings, parachutes, or spines that grab—reflect the dispersal strategies that help spread seeds.
Most modern angiosperms are classified as either monocots or eudicots, based on the structure of their leaves and embryos. Basal angiosperms, such as water lilies, are considered more primitive because they share morphological traits with both monocots and eudicots.
Flowers and Fruits as an Evolutionary Adaptation
Flowers: Angiosperms produce their gametes in separate organs, which are usually housed in a flower (Fig. 2). Both fertilization and embryo development take place inside an anatomical structure that provides a stable system of sexual reproduction largely sheltered from environmental fluctuations. Flowering plants are the most diverse phylum on Earth after insects; flowers come in a bewildering array of sizes, shapes, colors, smells, and arrangements. Most flowers have a mutualistic pollinator, with the distinctive features of flowers reflecting the nature of the pollination agent. The relationship between pollinator and flower characteristics is one of the great examples of coevolution.
Flowers are modified leaves, or sporophylls, organized around a central stalk. Although they vary greatly in appearance, all flowers contain the same structures: sepals, petals, carpels, and stamens. The peduncle attaches the flower to the plant. A whorl of sepals (collectively called the calyx) is located at the base of the peduncle and encloses the unopened floral bud. Sepals are usually photosynthetic organs, although there are some exceptions. For example, the corolla in lilies and tulips consists of three sepals and three petals that look virtually identical. Petals, collectively the corolla, are located inside the whorl of sepals and often display vivid colors to attract pollinators. Flowers pollinated by wind are usually small, feathery, and visually inconspicuous. Sepals and petals together form the perianth. The sexual organs (carpels and stamens) are located at the center of the flower.
Styles, stigmas, and ovules constitute the female organ: the gynoecium or carpel. The flower structure is very diverse, and carpels may be singular, multiple, or fused. Multiple fused carpels comprise a pistil. The megaspores and the female gametophytes are produced and protected by the thick tissues of the carpel. A long, thin structure called a style leads from the sticky stigma, where pollen is deposited, to the ovary, enclosed in the carpel. The ovary houses one or more ovules, each of which will develop into a seed upon fertilization. The male reproductive organs, the stamens (collectively called the androecium), surround the central carpel. Stamens are composed of a thin stalk called a filament and a sac-like structure called the anther. The filament supports the anther, where the microspores are produced by meiosis and develop into pollen grains.
Review Question:
Fruit: Following fertilization of the egg, the ovule grows into a seed (Fig. 3). The surrounding tissues of the ovary thicken, developing into a fruit that will protect the seed and often ensure its dispersal over a wide geographic range. Not all fruits develop from an ovary; such structures are “false fruits.” Like flowers, fruit can vary tremendously in appearance, size, smell, and taste. Tomatoes, walnut shells, and avocados are all examples of fruit. As with pollen and seeds, fruits also act as agents of dispersal. Some may be carried away by the wind. Many attract animals that will eat the fruit and pass the seeds through their digestive systems, then deposit the seeds in another location. Cockleburs are covered with stiff, hooked spines that can hook into fur (or clothing) and hitch a ride on an animal for long distances. The cockleburs that clung to the velvet trousers of an enterprising Swiss hiker, George de Mestral, inspired his invention of the loop and hook fastener he named Velcro.
As the seed develops, the walls of the ovary thicken and form the fruit. The seed forms in an ovary, which also enlarges as the seeds grow. In botany, a fertilized and fully grown, ripened ovary is a fruit. Many foods commonly called vegetables are actually fruit. Eggplants, zucchini, string beans, and bell peppers are all technically a fruit because they contain seeds and are derived from the thick ovary tissue. Acorns are nuts, and winged maple whirligigs (whose botanical name is samara) are also fruit. Botanists classify fruit into more than two dozen different categories, only a few of which are actually fleshy and sweet.
Mature fruit can be fleshy or dry. Fleshy fruit includes familiar berries, peaches, apples, grapes, and tomatoes. Rice, wheat, and nuts are examples of dry fruit. Another distinction is that not all fruits are derived from the ovary. For instance, strawberries are derived from the receptacle and apples from the pericarp or hypanthium. Some fruits are derived from separate ovaries in a single flower, such as the raspberry. Other fruits, such as the pineapple, form from clusters of flowers. Additionally, some fruits, like watermelon and orange, have rinds. Regardless of how they are formed, fruits are an agent of seed dispersal. The variety of shapes and characteristics reflect the mode of dispersal. The wind carries the light dry fruit of trees and dandelions. Water transports floating coconuts. Some fruits attract herbivores with color or perfume, or as food. Once eaten, tough, undigested seeds are dispersed through the herbivore’s feces. Other fruits have burs and hooks to cling to fur and hitch rides on animals.
The Life Cycle of an Angiosperm
The adult, or sporophyte, phase is the main phase of an angiosperm’s life cycle (Fig. 4). Like gymnosperms, angiosperms are heterosporous. Therefore, they generate microspores, which will generate pollen grains as the male gametophytes, and megaspores, which will form an ovule that contains female gametophytes. Inside the anthers’ microsporangia, male gametophytes divide by meiosis to generate haploid microspores, which, in turn, undergo mitosis and give rise to pollen grains. Each pollen grain contains two cells: one generative cell that will divide into two sperm and a second cell that will become the pollen tube cell.
If a flower lacked a megasporangium, what type of gamete would not form? If the flower lacked a microsporangium, what type of gamete would not form? The ovule, sheltered within the ovary of the carpel, contains the megasporangium protected by two layers of integuments and the ovary wall. Within each megasporangium, a megasporocyte undergoes meiosis, generating four megaspores—three small and one large. Only the large megaspore survives; it produces the female gametophyte, referred to as the embryo sac. The megaspore divides three times to form an eight-cell stage. Four of these cells migrate to each pole of the embryo sac; two come to the equator, and will eventually fuse to form a 2n polar nucleus; the three cells away from the egg form antipodals, and the two cells closest to the egg become the synergids. The mature embryo sac contains one egg cell, two synergids or “helper” cells, three antipodal cells, and two polar nuclei in a central cell. When a pollen grain reaches the stigma, a pollen tube extends from the grain, grows down the style, and enters through the micropyle: an opening in the integuments of the ovule. The two sperm cells are deposited in the embryo sac. A double fertilization event then occurs. One sperm and the egg combine, forming a diploid zygote—the future embryo. The other sperm fuses with the 2n polar nuclei, forming a triploid cell that will develop into the endosperm, which is the tissue that serves as a food reserve. The zygote develops into an embryo with a radicle or small root, and one (monocot) or two (dicot) leaf-like organs called cotyledons. This difference in the number of embryonic leaves is the basis for the two major groups of angiosperms: the monocots and the eudicots. Seed food reserves are stored outside the embryo, in the form of complex carbohydrates, lipids or proteins. The cotyledons serve as conduits to transmit the broken-down food reserves from their storage site inside the seed to the developing embryo. The seed consists of a toughened layer of integuments forming the coat, the endosperm with food reserves, and at the center, the well-protected embryo. Most flowers are monoecious or bisexual, which means that they carry both stamens and carpels; only a few species self-pollinate. Monoecious flowers are also known as “perfect” flowers because they contain both types of sex organs. Both anatomical and environmental barriers promote cross-pollination mediated by a physical agent (wind or water), or an animal, such as an insect or bird. Cross-pollination increases genetic diversity in a species.
Review Question:
Diversity of Angiosperms
Angiosperms are classified in a single phylum: the Anthophyta. Modern angiosperms appear to be a monophyletic group, which means that they originate from a single ancestor. Flowering plants are divided into two major groups, according to the structure of the cotyledons, pollen grains, and other structures. Monocots include grasses and lilies, and eudicots or dicots form a polyphyletic group (Fig. 5). Basal angiosperms are a group of plants that are believed to have branched off before the separation into monocots and eudicots because they exhibit traits from both groups. They are categorized separately in many classification schemes.
Basal Angiosperms
The Magnoliidae are represented by the magnolias: tall trees bearing large, fragrant flowers that have many parts and are considered archaic. Laurel trees produce fragrant leaves and small, inconspicuous flowers. The Laurales grow mostly in warmer climates and are small trees and shrubs. Familiar plants in this group include the bay laurel, cinnamon, spicebush, and avocado tree. The Nymphaeales are comprised of the water lilies, lotus, and similar plants; all species thrive in freshwater biomes and have leaves that float on the water surface or grow underwater. Water lilies are particularly prized by gardeners and have graced ponds and pools for thousands of years. The Piperales are a group of herbs, shrubs, and small trees that grow in tropical climates. They have small flowers without petals that are tightly arranged in long spikes. Many species are the source of prized fragrance or spices, for example, the berries of Piper nigrum are the familiar black peppercorns that are used to flavor many dishes.
Monocots
Plants in the monocot group are primarily identified as such by the presence of a single cotyledon in the seedling. Other anatomical features shared by monocots include veins that run parallel to the length of the leaves, and flower parts that are arranged in a three- or six-fold symmetry. True woody tissue is rarely found in monocots. In palm trees, vascular and parenchyma tissues produced by the primary and secondary thickening meristems form the trunk. The pollen from the first angiosperms was monosulcate, containing a single furrow or pore through the outer layer. This feature is still seen in modern monocots. The vascular tissue of the stem is not arranged in any particular pattern. The root system is mostly adventitious and unusually positioned, with no major taproot. The monocots include familiar plants such as the true lilies (which are at the origin of their alternate name of Liliopsida), orchids, grasses, and palms. Many important crops are monocots, such as rice and other cereals, corn, sugar cane, and tropical fruits like bananas and pineapples
Eudicots
Eudicots, or true dicots, are characterized by the presence of two cotyledons in the developing shoot. Veins form a network in leaves, and flower parts come in four, five, or many whorls. Vascular tissue forms a ring in the stem; in monocots, the vascular tissue is scattered in the stem. Eudicots can be herbaceous (like grasses) or produce woody tissues. Most eudicots produce pollen that is trisulcate or triporate, with three furrows or pores. The root system is usually anchored by one main root developed from the embryonic radicle. Eudicots comprise two-thirds of all flowering plants. Many species exhibit characteristics that belong to either group; as such, the classification of a plant as a monocot or a eudicot is not always clearly evident.
Review Question:
Summary
Angiosperms are the dominant form of plant life in most terrestrial ecosystems, comprising about 90 percent of all plant species. Most crops and ornamental plants are angiosperms. Their success comes from two innovative structures that protect reproduction from variability in the environment: the flower and the fruit. Flowers were derived from modified leaves. The main parts of a flower are the sepals and petals, which protect the reproductive parts: the stamens and the carpels. The stamens produce the male gametes in pollen grains. The carpels contain the female gametes (the eggs inside the ovules), which are within the ovary of a carpel. The walls of the ovary thicken after fertilization, ripening into fruit that ensures dispersal by wind, water, or animals.
The angiosperm life cycle is dominated by the sporophyte stage. Double fertilization is an event unique to angiosperms. One sperm in the pollen fertilizes the egg, forming a diploid zygote, while the other combines with the two polar nuclei, forming a triploid cell that develops into a food storage tissue called the endosperm. Flowering plants are divided into two main groups, the monocots and eudicots, according to the number of cotyledons in the seedlings. Basal angiosperms belong to an older lineage than monocots and eudicots.
End of Section Review Questions:
D) spore
Review: Floral Structures
Image Attributions
Figure 1. Image modified by D. Jennings. Origin unknown
Figure 2. Image courtesy of Mariana Ruiz LadyofHats [Public domain] Modified by D. Jennings
Figure 3. Image courtesy of CNX OpenStax [CC BY 4.0 (https://creativecommons.org/licenses/by/4.0)]
Figure 4. Image courtesy of LadyofHats Mariana Ruiz [Public domain]