Plant Diversity II: The Evolution of Seed Plants Chapter 30 Plant Diversity II: The Evolution of Seed Plants

Overview: Transforming the World Seeds changed the course of plant evolution, enabling their bearers to become the dominant producers in most terrestrial ecosystems Seed plants originated about 360 million years ago A seed consists of an embryo and nutrients surrounded by a protective coat Domestication of seed plants had begun by 8,000 years ago and allowed for permanent settlements © 2011 Pearson Education, Inc.

Figure 30.1 Figure 30.1 What human reproductive organ is functionally similar to this seed? 3

Concept 30.1: Seeds and pollen grains are key adaptations for life on land In addition to seeds, the following are common to all seed plants Reduced gametophytes Heterospory Ovules Pollen © 2011 Pearson Education, Inc.

Advantages of Reduced Gametophytes The gametophytes of seed plants develop within the walls of spores that are retained within tissues of the parent sporophyte © 2011 Pearson Education, Inc.

Ferns and other seedless Figure 30.2 PLANT GROUP Mosses and other nonvascular plants Ferns and other seedless vascular plants Seed plants (gymnosperms and angiosperms) Reduced, independent (photosynthetic and free-living) Reduced (usually microscopic), dependent on surrounding sporophyte tissue for nutrition Gametophyte Dominant Reduced, dependent on gametophyte for nutrition Sporophyte Dominant Dominant Gymnosperm Angiosperm Sporophyte (2n) Microscopic female gametophytes (n) inside ovulate cone Sporophyte (2n) Microscopic female gametophytes (n) inside these parts of flowers Gametophyte (n) Example Figure 30.2 Gametophyte-sporophyte relationships in different plant groups. Microscopic male gametophytes (n) inside these parts of flowers Microscopic male gametophytes (n) inside pollen cone Sporophyte (2n) Sporophyte (2n) Gametophyte (n) 6

Heterospory: The Rule Among Seed Plants The ancestors of seed plants were likely homosporous, while seed plants are heterosporous Megasporangia produce megaspores that give rise to female gametophytes Microsporangia produce microspores that give rise to male gametophytes © 2011 Pearson Education, Inc.

Ovules and Production of Eggs An ovule consists of a megasporangium, megaspore, and one or more protective integuments Gymnosperm megaspores have one integument Angiosperm megaspores usually have two integuments © 2011 Pearson Education, Inc.

(a) Unfertilized ovule Figure 30.3-1 Immature ovulate cone Integument (2n) Spore wall Megaspore (n) Figure 30.3 From ovule to seed in a gymnosperm. Megasporangium (2n) Micropyle Pollen grain (n) (a) Unfertilized ovule 9

Pollen and Production of Sperm Microspores develop into pollen grains, which contain the male gametophytes Pollination is the transfer of pollen to the part of a seed plant containing the ovules Pollen eliminates the need for a film of water and can be dispersed great distances by air or animals If a pollen grain germinates, it gives rise to a pollen tube that discharges sperm into the female gametophyte within the ovule © 2011 Pearson Education, Inc.

(a) Unfertilized ovule (b) Fertilized ovule Figure 30.3-2 Immature ovulate cone Female gametophyte (n) Integument (2n) Spore wall Egg nucleus (n) Megaspore (n) Discharged sperm nucleus (n) Figure 30.3 From ovule to seed in a gymnosperm. Megasporangium (2n) Pollen tube Micropyle Male gametophyte (n) Pollen grain (n) (a) Unfertilized ovule (b) Fertilized ovule 11

The Evolutionary Advantage of Seeds A seed develops from the whole ovule A seed is a sporophyte embryo, along with its food supply, packaged in a protective coat © 2011 Pearson Education, Inc.

(a) Unfertilized ovule (b) Fertilized ovule (c) Gymnosperm seed Figure 30.3-3 Immature ovulate cone Seed coat Female gametophyte (n) Integument (2n) Spore wall Spore wall Egg nucleus (n) Megaspore (n) Discharged sperm nucleus (n) Food supply (n) Figure 30.3 From ovule to seed in a gymnosperm. Megasporangium (2n) Pollen tube Male gametophyte (n) Embryo (2n) Micropyle Pollen grain (n) (a) Unfertilized ovule (b) Fertilized ovule (c) Gymnosperm seed 13

Seeds provide some evolutionary advantages over spores They may remain dormant for days to years, until conditions are favorable for germination Seeds have a supply of stored food They may be transported long distances by wind or animals © 2011 Pearson Education, Inc.

Concept 30.2: Gymnosperms bear “naked” seeds, typically on cones Gymnosperms means “naked seeds” The seeds are exposed on sporophylls that form cones Angiosperm seeds are found in fruits, which are mature ovaries © 2011 Pearson Education, Inc.

Nonvascular plants (bryophytes) Figure 30.UN01 Nonvascular plants (bryophytes) Seedless vascular plants Gymnosperms Angiosperms Figure 30.UN01 In-text figure, p. 621 16

Gymnosperm Evolution Fossil evidence reveals that by the late Devonian period some plants, called progymnosperms, had begun to acquire some adaptations that characterize seed plants © 2011 Pearson Education, Inc.

Figure 30.4 Figure 30.4 A progymnosperm. 18

Living seed plants can be divided into two clades: gymnosperms and angiosperms Gymnosperms appear early in the fossil record about 305 million years ago and dominated Mesozoic (251–65 million years ago) terrestrial ecosystems Gymnosperms were better suited than nonvascular plants to drier conditions © 2011 Pearson Education, Inc.

Angiosperms began to replace gymnosperms near the end of the Mesozoic Angiosperms now dominate more terrestrial ecosystems Today, cone-bearing gymnosperms called conifers dominate in the northern latitudes © 2011 Pearson Education, Inc.

The gymnosperm consist of four phyla Cycadophyta (cycads) Gingkophyta (one living species: Ginkgo biloba) Gnetophyta (three genera: Gnetum, Ephedra, Welwitschia) Coniferophyta (conifers, such as pine, fir, and redwood) © 2011 Pearson Education, Inc.

Phylum Cycadophyta Individuals have large cones and palmlike leaves These thrived during the Mesozoic, but relatively few species exist today © 2011 Pearson Education, Inc.

Cycas revoluta Figure 30.5a Figure 30.5 Exploring: Gymnosperm Diversity Cycas revoluta 23

Phylum Ginkgophyta This phylum consists of a single living species, Ginkgo biloba It has a high tolerance to air pollution and is a popular ornamental tree © 2011 Pearson Education, Inc.

Ginkgo biloba pollen-producing tree Figure 30.5b Figure 30.5 Exploring: Gymnosperm Diversity Ginkgo biloba leaves and fleshy seeds Ginkgo biloba pollen-producing tree 25

Phylum Gnetophyta This phylum comprises three genera Species vary in appearance, and some are tropical whereas others live in deserts © 2011 Pearson Education, Inc.

Ovulate cones Gnetum Welwitschia Ephedra Figure 30.5d Figure 30.5 Exploring: Gymnosperm Diversity Welwitschia Ephedra 27

Phylum Coniferophyta This phylum is by far the largest of the gymnosperm phyla Most conifers are evergreens and can carry out photosynthesis year round © 2011 Pearson Education, Inc.

Figure 30.5 Exploring: Gymnosperm Diversity Common juniper Douglas fir Figure 30.5 Exploring: Gymnosperm Diversity Sequoia European larch Wollemi pine Bristlecone pine 29

The Life Cycle of a Pine: A Closer Look Three key features of the gymnosperm life cycle are Dominance of the sporophyte generation Development of seeds from fertilized ovules The transfer of sperm to ovules by pollen The life cycle of a pine provides an example Animation: Pine Life Cycle © 2011 Pearson Education, Inc.

It takes nearly three years from cone production to mature seed The pine tree is the sporophyte and produces sporangia in male and female cones Small cones produce microspores called pollen grains, each of which contains a male gametophyte The familiar larger cones contain ovules, which produce megaspores that develop into female gametophytes It takes nearly three years from cone production to mature seed © 2011 Pearson Education, Inc.

Key Haploid (n) Diploid (2n) Pollen cone Microsporocytes Mature (2n) Figure 30.6-1 Key Haploid (n) Diploid (2n) Pollen cone Microsporocytes (2n) Mature sporophyte (2n) Pollen grains (n) MEIOSIS Microsporangia Microsporangium (2n) Figure 30.6 The life cycle of a pine. 32

Key Haploid (n) Ovule Diploid (2n) Ovulate Megasporocyte (2n) cone Figure 30.6-2 Key Haploid (n) Diploid (2n) Ovule Ovulate cone Megasporocyte (2n) Pollen cone Integument Microsporocytes (2n) Mature sporophyte (2n) Megasporangium (2n) Pollen grain Pollen grains (n) MEIOSIS MEIOSIS Microsporangia Microsporangium (2n) Surviving megaspore (n) Figure 30.6 The life cycle of a pine. 33

Key Haploid (n) Ovule Diploid (2n) Ovulate Megasporocyte (2n) cone Figure 30.6-3 Key Haploid (n) Diploid (2n) Ovule Ovulate cone Megasporocyte (2n) Pollen cone Integument Microsporocytes (2n) Mature sporophyte (2n) Megasporangium (2n) Pollen grain Pollen grains (n) MEIOSIS MEIOSIS Microsporangia Microsporangium (2n) Surviving megaspore (n) Archegonium Figure 30.6 The life cycle of a pine. Female gametophyte Sperm nucleus (n) Egg nucleus (n) Pollen tube FERTILIZATION 34

Pollen grain Food reserves (n) Figure 30.6-4 Key Haploid (n) Diploid (2n) Ovule Ovulate cone Megasporocyte (2n) Pollen cone Integument Microsporocytes (2n) Mature sporophyte (2n) Megasporangium (2n) Pollen grain Pollen grains (n) MEIOSIS MEIOSIS Microsporangia Microsporangium (2n) Surviving megaspore (n) Seedling Archegonium Figure 30.6 The life cycle of a pine. Female gametophyte Seeds Food reserves (n) Sperm nucleus (n) Egg nucleus (n) Seed coat (2n) Pollen tube Embryo (new sporophyte) (2n) FERTILIZATION 35

Concept 30.3: The reproductive adaptations of angiosperms include flowers and fruits Angiosperms are seed plants with reproductive structures called flowers and fruits They are the most widespread and diverse of all plants © 2011 Pearson Education, Inc.

Nonvascular plants (bryophytes) Figure 30.UN02 Nonvascular plants (bryophytes) Seedless vascular plants Gymnosperms Angiosperms Figure 30.UN02 In-text figure, p. 625 37

Characteristics of Angiosperms All angiosperms are classified in a single phylum, Anthophyta, from the Greek anthos for flower Angiosperms have two key adaptations Flowers Fruits © 2011 Pearson Education, Inc.

Flowers The flower is an angiosperm structure specialized for sexual reproduction Many species are pollinated by insects or animals, while some species are wind-pollinated © 2011 Pearson Education, Inc.

A flower is a specialized shoot with up to four types of modified leaves Sepals, which enclose the flower Petals, which are brightly colored and attract pollinators Stamens, which produce pollen Carpels, which produce ovules © 2011 Pearson Education, Inc.

A stamen consists of a stalk called a filament, with a sac called an anther where the pollen is produced A carpel consists of an ovary at the base and a style leading up to a stigma, where pollen is received Video: Flower Blooming (time lapse) © 2011 Pearson Education, Inc.

Stigma Carpel Stamen Anther Style Filament Ovary Petal Sepal Ovule Figure 30.7 Stigma Carpel Stamen Anther Style Filament Ovary Figure 30.7 The structure of an idealized flower. Petal Sepal Ovule 42

Fruits A fruit typically consists of a mature ovary but can also include other flower parts Fruits protect seeds and aid in their dispersal Mature fruits can be either fleshy or dry Animation: Fruit Development © 2011 Pearson Education, Inc.

Tomato Ruby grapefruit Nectarine Hazelnut Milkweed Figure 30.8 Figure 30.8 Some variations in fruit structure. Milkweed 44

Various fruit adaptations help disperse seeds Seeds can be carried by wind, water, or animals to new locations © 2011 Pearson Education, Inc.

Wings Seeds within berries Barbs Figure 30.9 Figure 30.9 Fruit adaptations that enhance seed dispersal. Barbs 46

The Angiosperm Life Cycle The flower of the sporophyte is composed of both male and female structures Male gametophytes are contained within pollen grains produced by the microsporangia of anthers The female gametophyte, or embryo sac, develops within an ovule contained within an ovary at the base of a stigma Most flowers have mechanisms to ensure cross-pollination between flowers from different plants of the same species © 2011 Pearson Education, Inc.

The ovule is entered by a pore called the micropyle A pollen grain that has landed on a stigma germinates and the pollen tube of the male gametophyte grows down to the ovary The ovule is entered by a pore called the micropyle Double fertilization occurs when the pollen tube discharges two sperm into the female gametophyte within an ovule © 2011 Pearson Education, Inc.

The triploid endosperm nourishes the developing embryo One sperm fertilizes the egg, while the other combines with two nuclei in the central cell of the female gametophyte and initiates development of food-storing endosperm The triploid endosperm nourishes the developing embryo Within a seed, the embryo consists of a root and two seed leaves called cotyledons © 2011 Pearson Education, Inc.

Microsporangium Anther Microsporocytes (2n) Mature flower on Figure 30.10-1 Microsporangium Anther Mature flower on sporophyte plant (2n) Microsporocytes (2n) MEIOSIS Microspore (n) Generative cell Tube cell Pollen grains Figure 30.10 The life cycle of an angiosperm. Key Haploid (n) Diploid (2n) 50

Microsporangium Anther Microsporocytes (2n) Mature flower on Figure 30.10-2 Microsporangium Anther Mature flower on sporophyte plant (2n) Microsporocytes (2n) MEIOSIS Microspore (n) Generative cell Ovule (2n) Male gametophyte (in pollen grain) (n) Tube cell Ovary Pollen grains MEIOSIS Megasporangium (2n) Surviving megaspore (n) Antipodal cells Female gametophyte (embryo sac) Central cell Synergids Pollen tube Figure 30.10 The life cycle of an angiosperm. Egg (n) Sperm (n) Key Haploid (n) Diploid (2n) 51

Discharged sperm nuclei (n) Figure 30.10-3 Microsporangium Anther Mature flower on sporophyte plant (2n) Microsporocytes (2n) MEIOSIS Microspore (n) Generative cell Ovule (2n) Male gametophyte (in pollen grain) (n) Tube cell Ovary Pollen grains MEIOSIS Stigma Megasporangium (2n) Pollen tube Sperm Surviving megaspore (n) Seed Antipodal cells Female gametophyte (embryo sac) Style Central cell Figure 30.10 The life cycle of an angiosperm. Synergids Pollen tube Egg (n) Sperm (n) Egg nucleus (n) FERTILIZATION Key Haploid (n) Diploid (2n) Discharged sperm nuclei (n) 52

Discharged sperm nuclei (n) Figure 30.10-4 Microsporangium Anther Mature flower on sporophyte plant (2n) Microsporocytes (2n) MEIOSIS Microspore (n) Generative cell Ovule (2n) Male gametophyte (in pollen grain) (n) Tube cell Ovary Germinating seed Pollen grains MEIOSIS Stigma Megasporangium (2n) Pollen tube Embryo (2n) Sperm Surviving megaspore (n) Endosperm (3n) Seed Seed coat (2n) Antipodal cells Female gametophyte (embryo sac) Style Central cell Pollen tube Figure 30.10 The life cycle of an angiosperm. Synergids Nucleus of developing endosperm (3n) Egg (n) Sperm (n) Egg nucleus (n) Zygote (2n) FERTILIZATION Key Haploid (n) Diploid (2n) Discharged sperm nuclei (n) 53

Angiosperm Evolution Darwin called the origin of angiosperms an “abominable mystery” Angiosperms originated at least 140 million years ago During the late Mesozoic, the major branches of the clade diverged from their common ancestor Scientists are studying fossils, refining phylogenies, and investigating developmental patterns to resolve the mystery © 2011 Pearson Education, Inc.

Fossil Angiosperms Chinese fossils of 125-million-year-old angiosperms share some traits with living angiosperms but lack others Archaefructus sinensis, for example, has anthers and seeds but lacks petals and sepals © 2011 Pearson Education, Inc.

(a) Archaefructus sinensis, a 125- million-year-old fossil Figure 30.11 Carpel Stamen 5 cm (a) Archaefructus sinensis, a 125- million-year-old fossil Figure 30.11 An early flowering plant. (b) Artist’s reconstruction of Archaefructus sinensis 56

Angiosperm Phylogeny The ancestors of angiosperms and gymnosperms diverged about 305 million years ago Angiosperms may be closely related to Bennettitales, extinct seed plants with flowerlike structures Amborella and water lilies are likely descended from two of the most ancient angiosperm lineages © 2011 Pearson Education, Inc.

Figure 30.12 Angiosperm evolutionary history. Living gymnosperms Bennettitales Microsporangia (contain microspores) Amborella Water lilies Most recent common ancestor of all living angiosperms Star anise and relatives Magnoliids Monocots Ovules Figure 30.12 Angiosperm evolutionary history. Eudicots 300 250 200 150 100 50 Millions of years ago (a) A possible ancestor of the angiosperms? (b) Angiosperm phylogeny 58

(a) A possible ancestor of the angiosperms? Figure 30.12a Microsporangia (contain microspores) Figure 30.12 Angiosperm evolutionary history. Ovules (a) A possible ancestor of the angiosperms? 59

Most recent common ancestor of all living angiosperms Star anise Figure 30.12b Living gymnosperms Bennettitales Amborella Water lilies Most recent common ancestor of all living angiosperms Star anise and relatives Magnoliids Monocots Figure 30.12 Angiosperm evolutionary history. Eudicots 300 250 200 150 100 50 Millions of years ago (b) Angiosperm phylogeny 60

Developmental Patterns in Angiosperms Egg formation in the angiosperm Amborella resembles that of the gymnosperms In early angiosperms, the two integuments appear to originate separately Researchers are currently studying expression of flower development genes in gymnosperm and angiosperm species © 2011 Pearson Education, Inc.

Angiosperm Diversity Angiosperms comprise more than 250,000 living species Previously, angiosperms were divided into two main groups Monocots (one cotyledon) Dicots (two dicots) DNA studies suggest that monocots form a clade, but dicots are polyphyletic © 2011 Pearson Education, Inc.

The clade eudicot (“true” dicots) includes most dicots The rest of the former dicots form several small lineages Basal angiosperms are less derived and include the flowering plants belonging to the oldest lineages Magnoliids share some traits with basal angiosperms but evolved later © 2011 Pearson Education, Inc.

Basal Angiosperms Three small lineages constitute the basal angiosperms These include Amborella trichopoda, water lilies, and star anise © 2011 Pearson Education, Inc.

Basal Angiosperms Star anise Water lily Amborella trichopoda Figure 30.13a Basal Angiosperms Star anise Water lily Figure 30.13 Exploring: Angiosperm Diversity Amborella trichopoda 65

Magnoliids Magnoliids include magnolias, laurels, and black pepper plants Magnoliids are more closely related to monocots and eudicots than basal angiosperms © 2011 Pearson Education, Inc.

Magnoliids Southern magnolia Figure 30.13b Figure 30.13 Exploring: Angiosperm Diversity Southern magnolia 67

Monocots More than one-quarter of angiosperm species are monocots © 2011 Pearson Education, Inc.

Monocots Orchid Lily Pygmy date palm Anther Stigma Ovary Filament Figure 30.13c Monocots Orchid Lily Figure 30.13 Exploring: Angiosperm Diversity Pygmy date palm Anther Stigma Ovary Filament Barley, a grass 69

Eudicots More than two-thirds of angiosperm species are eudicots © 2011 Pearson Education, Inc.

Eudicots Dog rose California poppy Pyrenean oak Snow pea Zucchini Figure 30.13d Eudicots Dog rose California poppy Figure 30.13 Exploring: Angiosperm Diversity Pyrenean oak Snow pea Zucchini 71

Figure 30.13 Exploring: Angiosperm Diversity Monocot Characteristics Eudicot Characteristics Embryos One cotyledon Two cotyledons Leaf venation Veins usually parallel Veins usually netlike Stems Vascular tissue usually arranged in ring Vascular tissue scattered Roots Root system usually fibrous (no main root) Taproot (main root) usually present Figure 30.13 Exploring: Angiosperm Diversity Pollen Pollen grain with one opening Pollen grain with three openings Flowers Floral organs usually in multiples of three Floral organs usually in multiples of four or five 72

Evolutionary Links Between Angiosperms and Animals Animals influence the evolution of plants and vice versa For example, animal herbivory selects for plant defenses For example, interactions between pollinators and flowering plants select for mutually beneficial adaptations Video: Bat Pollinating Agave Plant Video: Bee Pollinating © 2011 Pearson Education, Inc.

Figure 30.14 Figure 30.14 A plant pollinated by flies. 74

Clades with bilaterally symmetrical flowers have more species than those with radially symmetrical flowers This is likely because bilateral symmetry affects the movement of pollinators and reduces gene flow in diverging populations © 2011 Pearson Education, Inc.

Figure 30.15 Figure 30.15 Pollinating a bilaterally symmetrical flower. 76

Concept 30.4: Human welfare depends greatly on seed plants No group of plants is more important to human survival than seed plants Plants are key sources of food, fuel, wood products, and medicine Our reliance on seed plants makes preservation of plant diversity critical © 2011 Pearson Education, Inc.

Products from Seed Plants Most of our food comes from angiosperms Six crops (wheat, rice, maize, potatoes, cassava, and sweet potatoes) yield 80% of the calories consumed by humans Modern crops are products of relatively recent genetic change resulting from artificial selection Many seed plants provide wood Secondary compounds of seed plants are used in medicines © 2011 Pearson Education, Inc.

Table 30.1 Table 30.1 Examples of Plant-Derived Medicines 79

Threats to Plant Diversity Destruction of habitat is causing extinction of many plant species In the tropics 55,000 km2 are cleared each year At this rate, the remaining tropical forests will be eliminated in 200 years Loss of plant habitat is often accompanied by loss of the animal species that plants support © 2011 Pearson Education, Inc.

The tropical rain forests may contain undiscovered medicinal compounds At the current rate of habitat loss, 50% of Earth’s species will become extinct within the next 100–200 years The tropical rain forests may contain undiscovered medicinal compounds © 2011 Pearson Education, Inc.

A satellite image from 2000 shows clear-cut areas in Brazil surrounded Figure 30.16 A satellite image from 2000 shows clear-cut areas in Brazil surrounded by dense tropical forest. By 2009, much more of this same tropical forest had been cut down. Figure 30.16 Impact: Clear-Cutting of Tropical Forests 4 km 82