2Plants without Seeds: From Sea to Land The Plant KingdomThe Conquest of the LandThe Nontracheophytes: Liverworts, Hornworts, and MossesIntroducing the TracheophytesThe Surviving Nonseed Tracheophytes
3The Plant KingdomA plant is a photosynthetic eukaryote that uses chlorophylls a and b, stores carbohydrates, and develops from an embryo protected by tissues of the parent plant.Because of their development from embryos, plants are sometimes referred to as embryophytes.The kingdom Plantae is monophyletic, forming a single branch of the evolutionary tree.
5The Plant KingdomThe surviving members of the kingdom Plantae fall naturally into ten phyla.The seven plant phyla whose members possess well-developed vascular systems are called the tracheophytes.The remaining three phyla (liverworts, hornworts, and mosses) lack tracheids and are collectively referred to as the nontracheophytes.
8The Plant KingdomAlternation of generations is a universal feature of the life cycles of plants.This life cycle includes both multicellular diploid individuals and multicellular haploid individuals.Gametes are produced by mitosis, not meiosis. Meiosis produces spores that develop into multicellular haploid individuals.
9The Plant KingdomThe multicellular, diploid plant is called the sporophyte.Cells contained in the sporangia on the sporophyte produce haploid, unicellular spores by meiosis.The multicellular, haploid plant formed by mitosis and cytokinesis of a spore is called the gametophyte.The gametophyte produces haploid gametes.The fusion of two gametes results in the formation of a diploid cell, the zygote, and the cycle repeats.
11The Plant KingdomThe sporophyte generation extends from the zygote through the adult, multicellular, diploid plant.The gametophyte generation extends from the spore through the adult, multicellular, haploid plant to the gamete.Some protist life cycles also feature alternation of generations, suggesting that the plants arose from one of these protist groups.
12The Plant KingdomEvidence indicates that the closest living relatives of the plants are a group of green algae called charophytes.It has not been determined which charophyte clade (stoneworts or Coleochaete) is the true sister group to the plants.The ancestral green algae lived at the margins of ponds or marshes. From these marginal habitats, early plants made the move onto land.
13Figure 29.3 The Closest Relatives of Land Plants
14The Conquest of the Land Plants or their immediate ancestors pioneered and modified the terrestrial environment, first invading 400–500 mya.The availability of water is a key difference between the aquatic and terrestrial environments.Land plants had to adapt to new challenges in the terrestrial environment, such as obtaining water, dealing with gravity, and dispersing gametes.
15The Conquest of the Land Some adaptations to life on land:The cuticle, a waxy covering that prevents dryingGametangia, cases that enclose gametes and prevent dryingEmbryos, young sporophytes contained within a protective structurePigments that afford protection against mutagenic ultraviolet radiationThick spore walls to prevent drying and resist decayA mutualistic association with a fungus that promotes nutrient uptake from the soil
16The Conquest of the Land The first plants lacked vascular tissue.Some of the mosses, which are nontracheophytes, have a small amount of simple conducting tissue.The true tracheophytes, which have specialized conducting cells called tracheids, arose later.
18The Conquest of the Land The nontracheophytes have developed ways to obtain water and minerals in the absence of a vascular system:Many grow in dense masses through which water can move by capillary action.They have leaflike structures that catch and hold water that splashes onto them.They are small enough that minerals can be distributed evenly by diffusion.
19The Conquest of the Land The tracheophytes have a well-developed vascular system which consists of two specialized tissues:Phloem conducts products of photosynthesis from sites where they are produced to sites where they are used or stored.Xylem conducts water and minerals from the soil to the aerial parts of the plants. Xylem, stiffened by a substance called lignin, also provides support in the terrestrial environment.
20The Nontracheophytes: Liverworts, Hornworts, and Mosses The nontracheophytes usually grow in dense mats in moist habitats and are generally small in size.Layers of maternal tissue prevent loss of water from the embryo.The nontracheophytes have a thin cuticle, though it is not highly effective in retarding water loss.The nontracheophytes are widespread across six continents.
21The Nontracheophytes: Liverworts, Hornworts, and Mosses In nontracheophytes, the familiar green structure visible to the naked eye is the gametophyte.A nontracheophyte sporophyte produces unicellular, haploid spores as products of meiosis within a sporangium or capsule.The spore germinates and gives rise to a multicellular, haploid gametophyte whose cells contain chloroplasts.
23The Nontracheophytes: Liverworts, Hornworts, and Mosses Gametangia are specialized sex organs where gametes are formed.The archegonium is a multicellular, flask-shaped female sex organ with a long neck and a swollen base that contains a single egg.The antheridium is a male sex organ in which sperm are produced in large numbers.The sporophyte produces a sporangium, or capsule, within which meiotic divisions produce spores and thus the next gametophyte generation.
25The Nontracheophytes: Liverworts, Hornworts, and Mosses Liverworts, phylum Hepatophyta, may be the most ancient surviving plant clade.Rhizoids are water-absorbing filaments that are found on the lower surfaces of the simplest liverwort gametophytes.Liverwort sporophytes have a stalk that connects the capsule and the foot. This capsule can elongate to raise the capsule above ground level, aiding in the dispersion of spores.Other liverworts utilize springlike structures to disseminate their spores.
26The Nontracheophytes: Liverworts, Hornworts, and Mosses The hornworts, phylum Anthocerophyta, along with the mosses and the tracheophytes, all have an adaptation to life on land not found in the liverworts.These groups all possess stomata that allow the uptake of CO2 and the release of O2, but they can be closed to prevent excessive water loss.
28The Nontracheophytes: Liverworts, Hornworts, and Mosses Two characteristics distinguish hornworts from liverworts and mosses:The cells of hornworts contain a single large, platelike chloroplast, whereas liverworts and mosses contain numerous small, lens-shaped chloroplasts.Unlike the moss or liverwort sporophyte, whose stalk stops growing as the capsule matures, the hornwort sporophyte has no stalk.
29The Nontracheophytes: Liverworts, Hornworts, and Mosses Cyanobacteria often populate internal, mucilage- filled cavities within hornworts.These cyanobacteria are able to fix atmospheric nitrogen gas into a form that can be used by the hornwort.The exact evolutionary status of hornworts is still unresolved.
30The Nontracheophytes: Liverworts, Hornworts, and Mosses The phylum Bryophyta (mosses) are probably sister to the tracheophytes.Hydroid cells, found in many mosses, are a likely progenitor of the water-conducting cells of the tracheophytes.When hydroid cells die, they leave a tiny channel through which water can flow.The sporophytes of mosses and tracheophytes grow by apical cell division, whereby a region at the growing tip provides an organized pattern of cell division, elongation, and differentiation.
32The Nontracheophytes: Liverworts, Hornworts, and Mosses Sporophyte development results in the formation of an absorptive foot anchored to the gametophyte, a stalk, and a capsule.After meiosis and spore development are complete, the top of the capsule is shed.A series of toothlike structures surrounds the opening of the capsule and digs into the mass of spores when the atmosphere is dry.When the atmosphere becomes moist, they are flung out. Thus the spores are dispersed when conditions favor germination.
34Introducing the Tracheophytes Although the tracheophytes are a large and diverse group, their appearance can be attributed to a single evolutionary event.The sporophyte generation of a now-extinct organism produced a new cell type, called the tracheid.The tracheid is the principal water-conducting element in the xylem in all tracheophytes except the angiosperms.
35Introducing the Tracheophytes There are seven distinct phyla that are present- day evolutionary descendants of the early tracheophytes.These seven phyla can be sorted into two groups: those that produce seeds and those that do not.
37Introducing the Tracheophytes The plant kingdom invaded land between 400 and 500 million years ago.During the Devonian period the appearance and proliferation of the club mosses (lycopods), horsetails, and ferns made the environment more hospitable to animals.Trees became dominant during the Carboniferous period. The tropical swamp forests would become coal deposits.At the end of the Permian period, the 200-million- year reign of the lycopod–fern forests came to an end as they were replaced by forests of seed plants.
39Introducing the Tracheophytes The first tracheophytes belonged to the now- extinct phylum Rhyniophyta.The rhyniophytes had early versions of the structural features found in all other tracheophyte phyla.
41Introducing the Tracheophytes The first rhyniophytes were found in Devonian rocks near Rhynie, Scotland.The fossil plants had a simple vascular system of xylem and phloem. They lacked leaves and roots but were anchored to the soil by horizontal portions of stem called rhizomes.Inspection of fossil sporangia showed that the spores were in groups of four.Most living nonseed tracheophytes show an arrangement of spores such as this only in the sporophyte immediately after meiosis. It was concluded that the Rhynie fossils must be sporophytes.
42Introducing the Tracheophytes The phylum Lycophyta, the club mosses, appeared in the Silurian period.The phylum Pteridophyta, the ferns, horsetails, and whisk ferns, appeared in the Devonian period.These new groups had true roots, true leaves, and a differentiation between two types of spores.
44Introducing the Tracheophytes Plants that bear two distinct types of spores evolved later, and are said to be heterosporous.In heterosporous plants, the megaspore develops into a larger, specifically female gametophyte (megagametophyte).The microspore develops into the smaller, male gametophyte (microgametophyte).Heterospory evolved independently and repeatedly, suggesting that it affords selective advantages.
46The Surviving Nonseed Tracheophytes The nonseed tracheophytes have a large, independent sporophyte and a small, independent, short-lived gametophyte.The single-celled spore is the the most prominent resting stage of the life cycle.Nonseed tracheophytes must have an aqueous environment in order to be fertilized by the motile, flagellated sperm.Today, the ferns are the most abundant and diverse phylum of the nonseed tracheophytes.
47The Surviving Nonseed Tracheophytes The club mosses (phylum Lycophyta) have microphylls, exhibit apical growth, and have roots that branch dichotomously.Sporangia in many club mosses are contained within conelike structures called strobili, clusters of spore-bearing leaves inserted between a specialized leaf and the stem.There are both homosporous and heterosporous species.The Lycophyta and the Pteridophyta were the dominant phyla during the Carboniferous period.
49The Surviving Nonseed Tracheophytes The horsetails, whisk ferns, and ferns form a clade, the phylum Pteridophyta.The horsetails (all are genus Equisetum) have true roots that branch irregularly, and sporangia on short stalks called sporangiophores.The leaves are reduced megaphylls and grow in whorls.Stem growth is from the base of the stem segments.
51The Surviving Nonseed Tracheophytes The whisk ferns are two genera of rootless, spore- bearing plants, Psilotum and Tmesipteris.Psilotum has only minute scales instead of true leaves.Although whisk ferns resemble the most ancient tracheophytes, they are now considered to be highly specialized plants that evolved fairly recently.
53The Surviving Nonseed Tracheophytes The sporophytes of the ferns typically have true roots, stems, and leaves.The ferns first appeared during the Devonian period.About 97 percent of fern species belong to one clade, the leptosporangiate ferns. These ferns have sporangia with walls only one cell thick, borne on a stalk.Ferns are characterized by fronds, large leaves with complex vasculature.Sporangia are found on the undersurfaces of the fronds; in most species, they are clustered in groups called sori.
54Figure 29.19 Fern Sori Are Clusters of Sporangia
56The Surviving Nonseed Tracheophytes The sporophyte generation dominates the fern life cycle.A spore germinates and forms a heart-shaped gametophyte, bearing antheridia or archegonia (or both) on its underside.The antheridia release sperm that swim to a nearby archegonium and fertilize an egg.The sperm are guided by chemical attractants released from the archegonia.The resulting diploid embryo forms roots and fronds, and grows into the familiar sporophyte life stage.