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Figure 38.1 CAMPBELL BIOLOGY Figure 38.1a Flowers of Deceit TENTH EDITION Reece • Urry • Cain • Wasserman • Minorsky • Jackson ! Insects help angiosperms to reproduce sexually with physically distant members of their own species 38 ! For example, male long-horned bees mistake Ophrys flowers for females and attempt to mate with them Angiosperm Reproduction and Biotechnology ! The flower is pollinated in the process ! Unusually, the flower does not produce nectar and the male receives no benefit Lecture Presentation by Nicole Tunbridge and Kathleen Fitzpatrick 2 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Concept 38.1: Flowers, double fertilization, and fruits are key features of the angiosperm life cycle ! Many angiosperms lure insects with nectar; both plant and pollinator benefit ! Plant life cycles are characterized by the alternation between sporophyte (spore-producing) and gametophyte (gamete-producing) generations ! Mutualistic symbioses are common between plants and other species 3 © 2014 Pearson Education, Inc. ! Angiosperms can reproduce sexually and asexually 4 © 2014 Pearson Education, Inc. Flower Structure and Function ! In angiosperms, the sporophyte is the plant that we see; they are larger, more conspicuous and longer-lived than gametophytes ! Flowers are the reproductive shoots of the angiosperm sporophyte; they attach to a part of the stem called the receptacle ! The angiosperm life cycle is characterized by “three Fs”: f lowers, double fertilization, and f ruits ! Flowers consist of four floral organs: carpels, stamens, petals, and sepals ! Stamens and carpels are reproductive organs; sepals and petals are sterile ! Angiosperms are the most important group of plants in terrestrial ecosystems and in agriculture 5 © 2014 Pearson Education, Inc. 6 © 2014 Pearson Education, Inc. 7 © 2014 Pearson Education, Inc. 8 © 2014 Pearson Education, Inc. Figure 38.2 Stigma Stamen Anther Video: Flower Blooming (Time Lapse) Carpel Style Filament ! A carpel has a long style with a stigma on which pollen may land Ovary ! Complete flowers contain all four floral organs ! At the base of the style is an ovary containing one or more ovules ! Incomplete flowers lack one or more floral organs, for example stamens or carpels ! Clusters of flowers are called inflorescences ! A single carpel or group of fused carpels is called a pistil Petal ! A stamen consists of a filament topped by an anther with pollen sacs that produce pollen Sepal Ovule Receptacle 9 © 2014 Pearson Education, Inc. 10 © 2014 Pearson Education, Inc. 11 © 2014 Pearson Education, Inc. Figure 38.3a 12 © 2014 Pearson Education, Inc. Figure 38.3aa Figure 38.3ab Bilateral symmetry ! Much of floral diversity represents adaptation to specific pollinators ! Four general trends can be seen in the evolution of flowers ! Bilateral symmetry Musk mallow (radial symmetry) ! Reduction in the number of floral parts Musk mallow (radial symmetry) ! Fusion of floral parts “Bramley” orchid (bilateral symmetry) ! Location of ovaries inside receptacles 13 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. “Bramley” orchid (bilateral symmetry) 14 15 © 2014 Pearson Education, Inc. 16 © 2014 Pearson Education, Inc. Figure 38.3b Figure 38.3ba Reduction in number of floral parts Figure 38.3bb Figure 38.3c Fusion of floral parts Bloodroot Bloodroot Star of Bethlehem Drooping trillium Hedge bindweed 17 Drooping trillium © 2014 Pearson Education, Inc. 18 © 2014 Pearson Education, Inc. Figure 38.3ca 19 © 2014 Pearson Education, Inc. Figure 38.3cb 20 © 2014 Pearson Education, Inc. Figure 38.3d Figure 38.3da Ovaries located inside receptacles Ovary Hedge bindweed Star of Bethlehem Ovary Stone plant (longitudinal section) Stone plant (longitudinal section) Ovary Japanese quince (longitudinal section) 21 © 2014 Pearson Education, Inc. 22 © 2014 Pearson Education, Inc. 23 © 2014 Pearson Education, Inc. Figure 38.3db Figure 38.4-1 Carpel The Angiosperm Life Cycle: An Overview 24 © 2014 Pearson Education, Inc. Mature flower on sporophyte plant (2n) ! The angiosperm life cycle includes Anther MEIOSIS Carpel Anther Generative cell Tube cell Tube nucleus MEIOSIS Ovary Ovule with MEIOSIS megasporangium (2n) Pollen grains ! Pollination ! Double fertilization Female gametophyte (embryo sac) ! Seed development Ovary Japanese quince (longitudinal section) Key 25 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Figure 38.4-3 Carpel Anther MEIOSIS Carpel Ovary Ovule with MEIOSIS megasporangium (2n) Female gametophyte (embryo sac) Antipodal cells Polar nuclei in central cell Synergids Egg (n) Male gametophyte (in pollen grain) (n) Generative cell Tube cell Tube nucleus Megasporangium (2n) Surviving megaspore (n) Integuments Micropyle Stigma Pollen tube Sperm Tube nucleus Female gametophyte (embryo sac) Nucleus of developing endosperm (3n) Egg nucleus (n) FERTILIZATION Key Haploid (n) Diploid (2n) © 2014 Pearson Education, Inc. MEIOSIS Embryo (2n) Endosperm (3n) Seed coat (2n) Style Figure 38.4b Seed Antipodal cells Polar nuclei in central cell Synergids Egg (n) Microspore (n) Male gametophyte (in pollen grain) (n) Generative cell Tube cell Tube nucleus Megasporangium (2n) Surviving megaspore (n) Integuments Micropyle Carpel Anther Pollen grains Microsporangium (pollen sac) Microsporocytes (2n) MEIOSIS Stigma Pollen tube Sperm Tube nucleus Style Key Haploid (n) Diploid (2n) Zygote (2n) Egg nucleus (n) Microspore (n) Male gametophyte (in pollen grain) (n) Generative cell Tube cell Tube nucleus Ovary Ovule with MEIOSIS megasporangium (2n) Key Haploid (n) Diploid (2n) FERTILIZATION Key Discharged sperm nuclei (n) 29 Haploid (n) Diploid (2n) © 2014 Pearson Education, Inc. 28 Microsporocytes (2n) Ovary Ovule with MEIOSIS megasporangium (2n) Germinating seed Pollen grains Pollen grains Megasporangium (2n) Surviving megaspore (n) Integuments Micropyle © 2014 Pearson Education, Inc. Figure 38.4a Microsporangium (pollen sac) Anther Mature flower on sporophyte plant (2n) Microspore (n) Generative cell Tube cell Tube nucleus Haploid (n) Diploid (2n) 27 © 2014 Pearson Education, Inc. Figure 38.4-4 Microsporangium (pollen sac) Microsporocytes (2n) Mature flower on sporophyte plant (2n) Antipodal cells Polar nuclei in central cell Synergids Egg (n) Microspore (n) Male gametophyte (in pollen grain) (n) Key Haploid (n) Diploid (2n) 26 Microsporangium (pollen sac) Microsporocytes (2n) Mature flower on sporophyte plant (2n) Microspore (n) Male gametophyte (in pollen grain) (n) ! Gametophyte development Figure 38.4-2 Microsporangium (pollen sac) Microsporocytes (2n) Discharged sperm nuclei (n) 30 31 © 2014 Pearson Education, Inc. Megasporangium (2n) Surviving megaspore (n) Integuments Micropyle 32 © 2014 Pearson Education, Inc. Figure 38.4c Figure 38.4d Pollen grains Antipodal cells Female gametophyte (embryo sac) Megasporangium (2n) Surviving megaspore (n) Integuments Micropyle Polar nuclei in central cell Synergids Egg (n) Female gametophyte (embryo sac) Tube nucleus Style Video: Flowering Plant Life Cycle (Time Lapse) Embryo (2n) Endosperm (3n) Seed Seed coat (2n) Stigma Pollen tube Sperm Nucleus of developing endosperm (3n) Animation: Plant Fertilization Antipodal cells Polar nuclei in central cell Synergids Egg (n) Style Zygote (2n) Egg nucleus (n) FERTILIZATION Key Key Haploid (n) Diploid (2n) 33 © 2014 Pearson Education, Inc. Haploid (n) Diploid (2n) Discharged sperm nuclei (n) 34 © 2014 Pearson Education, Inc. 35 © 2014 Pearson Education, Inc. 36 © 2014 Pearson Education, Inc. Gametophyte Development ! Angiosperm gametophytes are microscopic and their development is obscured by protective tissues Development of Female Gametophytes (Embryo Sacs) ! The megaspore divides without cytokinesis, producing one large cell with eight nuclei Development of Male Gametophytes in Pollen Grains ! The embryo sac, or female gametophyte, develops within the ovule ! This cell is partitioned into a multicellular female gametophyte, the embryo sac ! Pollen develops from microspores within the microsporangia, or pollen sacs, of anthers ! Within an ovule, two integuments surround a megasporangium ! Each microspore undergoes mitosis to produce two cells: the generative cell and the tube cell ! One cell in the megasporangium undergoes meiosis, producing four megaspores, only one of which survives ! A pollen grain consists of the two-celled male gametophyte and the spore wall 37 38 39 40 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Pollination Double Fertilization Seed Development Methods of Pollination ! In angiosperms, pollination is the transfer of pollen from an anther to a stigma ! Fertilization, the fusion of gametes, occurs after the two sperm reach the female gametophyte ! After double fertilization, each ovule develops into a seed ! The transfer of pollen from anthers to stigma can be accomplished by wind, water, or animals ! After landing on a receptive stigma, a pollen grain produces a pollen tube that grows down into the ovary and discharges two sperm cells near the embryo sac ! One sperm fertilizes the egg, and the other combines with the two polar nuclei, giving rise to the triploid food-storing endosperm (3n) ! The ovary develops into a fruit enclosing the seed ! Wind-pollinated species (e.g., grasses and many trees) release large amounts of pollen ! This double fertilization ensures that endosperm only develops in ovules containing fertilized eggs 41 © 2014 Pearson Education, Inc. ! When a seed germinates, the embryo develops into a new sporophyte 42 © 2014 Pearson Education, Inc. Figure 38.5a Figure 38.5aa Abiotic pollination by wind 43 © 2014 Pearson Education, Inc. 44 © 2014 Pearson Education, Inc. Figure 38.5ab Figure 38.5ac Pollination by bees Common dandelion Common dandelion under normal light under ultraviolet light Hazel carpellate flower (carpels only) Hazel carpellate flower (carpels only) Hazel staminate flowers (stamens only) releasing clouds of pollen 45 © 2014 Pearson Education, Inc. Common dandelion under normal light Hazel staminate flowers (stamens only) releasing clouds of pollen 46 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. 47 48 © 2014 Pearson Education, Inc. Figure 38.5ad Figure 38.5b Figure 38.5ba Pollination by bats Pollination by moths and butterflies Figure 38.5bb Pollination by flies Anther Moth Anther Moth Stigma Long-nosed bat feeding on cactus flower at night Moth on yucca flower Blowfly on carrion flower Pollination by birds Common dandelion under ultraviolet light Stigma Long-nosed bat feeding on cactus flower at night Moth on yucca flower 49 © 2014 Pearson Education, Inc. Hummingbird drinking nectar of columbine flower 50 © 2014 Pearson Education, Inc. Figure 38.5bc 51 52 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Video: Bat Pollinating Agave Plant Video: Bee Pollinating Figure 38.5bd Blowfly on carrion flower Hummingbird drinking nectar of columbine flower 53 © 2014 Pearson Education, Inc. 54 © 2014 Pearson Education, Inc. 55 56 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. From Seed to Flowering Plant: A Closer Look Endosperm Development Figure 38.6 ! Coevolution is the joint evolution of interacting species in response to selection imposed by each other ! The development of a seed into a flowering plant includes several stages ! Endosperm development usually precedes embryo development ! Endosperm development ! Many flowering plants have coevolved with specific pollinators ! In most monocots and many eudicots, endosperm stores nutrients that can be used by the seedling ! Embryo development ! The shapes and sizes of flowers often correspond to the pollen transporting parts of their animal pollinators ! Seed germination ! Seedling development ! For example, Darwin correctly predicted a moth with a 28-cm-long tongue based on the morphology of a particular flower 57 © 2014 Pearson Education, Inc. ! In other eudicots, the food reserves of the endosperm are exported to the cotyledons ! Seed dormancy ! Flowering 58 © 2014 Pearson Education, Inc. 59 60 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Animation: Seed Development Structure of the Mature Seed Figure 38.7 Embryo Development Ovule Endosperm nucleus Zygote ! The first mitotic division of the zygote splits the fertilized egg into a basal cell and a terminal cell ! The embryo and its food supply are enclosed by a hard, protective seed coat Integuments ! The basal cell produces a multicellular suspensor, which anchors the embryo to the parent plant ! The seed enters a state of dormancy Zygote Terminal cell Basal cell ! The terminal cell gives rise to most of the embryo ! A mature seed is only about 5–15% water Proembryo ! The cotyledons form and the embryo elongates Suspensor Basal cell Cotyledons Shoot apex Root apex 61 © 2014 Pearson Education, Inc. Suspensor © 2014 Pearson Education, Inc. Seed coat Endosperm 62 63 © 2014 Pearson Education, Inc. 64 © 2014 Pearson Education, Inc. Figure 38.8 Figure 38.8a Seed coat Figure 38.8b Epicotyl Hypocotyl Radicle ! In some eudicots, such as the common garden bean, the embryo consists of the embryonic axis attached to two fleshy cotyledons (seed leaves) Cotyledons Seed coat Endosperm Cotyledons Epicotyl Hypocotyl (a) Common garden bean, a eudicot with thick cotyledons ! Below the cotyledons the embryonic axis is called the hypocotyl and terminates in the radicle (embryonic root); above the cotyledons it is called the epicotyl Endosperm Cotyledons Epicotyl Hypocotyl Radicle Hypocotyl Cotyledons Radicle Radicle (b) Castor bean, a eudicot with thin cotyledons Scutellum (cotyledon) ! The plumule comprises the epicotyl, young leaves, and shoot apical meristem Coleoptile Coleorhiza 65 © 2014 Pearson Education, Inc. Epicotyl Seed coat Seed coat © 2014 Pearson Education, Inc. Pericarp fused with seed coat Endosperm Epicotyl Hypocotyl Radicle (a) Common garden bean, a eudicot with thick cotyledons 66 (c) Maize, a monocot (b) Castor bean, a eudicot with thin cotyledons 67 © 2014 Pearson Education, Inc. 68 © 2014 Pearson Education, Inc. Figure 38.8c Seed Dormancy: An Adaptation for Tough Times Scutellum (cotyledon) Coleoptile Coleorhiza ! The seeds of some eudicots, such as castor beans, have thin cotyledons Pericarp fused with seed coat Endosperm Epicotyl Hypocotyl Radicle ! A monocot embryo has one cotyledon ! Grasses, such as maize and wheat, have a special cotyledon called a scutellum ! Two sheathes enclose the embryo of a grass seed: a coleoptile covering the young shoot and a coleorhiza covering the young root (c) Maize, a monocot ! Seed dormancy increases the chances that germination will occur at a time and place most advantageous to the seedling ! The breaking of seed dormancy often requires environmental cues, such as temperature or lighting changes ! Most seeds remain viable after a year or two of dormancy, but some last only days and others can remain viable for centuries 69 © 2014 Pearson Education, Inc. 70 © 2014 Pearson Education, Inc. 71 © 2014 Pearson Education, Inc. 72 © 2014 Pearson Education, Inc. Figure 38.9 Figure 38.9a Foliage leaves Cotyledon Seed Germination and Seedling Development Hypocotyl ! Germination depends on imbibition, the uptake of water due to low water potential of the dry seed ! In many eudicots, a hook forms in the hypocotyl, and growth pushes the hook above ground ! The radicle (embryonic root) emerges first; the developing root system anchors the plant ! Light causes the hook to straighten and pull the cotyledons and shoot tip up Epicotyl Foliage leaves Cotyledon Cotyledon Cotyledon Hypocotyl Hypocotyl Hypocotyl Epicotyl Cotyledon Radicle Hypocotyl Seed coat (a) Common garden bean ! Next, the shoot tip breaks through the soil surface Cotyledon Hypocotyl Foliage leaves Coleoptile Radicle Coleoptile Seed coat (a) Common garden bean 73 74 (b) Maize © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Radicle 75 76 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Flowering Fruit Structure and Function Figure 38.9b ! In maize and other grasses, which are monocots, the coleoptile pushes up through the soil creating a tunnel for the shoot tip to grow through Foliage leaves Coleoptile (b) Maize Coleoptile ! A fruit is the mature ovary of a flower ! It protects the enclosed seeds and aids in seed dispersal by wind or animals ! Flowering is triggered by a combination of environmental cues and internal signals ! In some fruits, such as soybean pods, the ovary wall dries out at maturity, whereas in other fruits, such as grapes, it remains fleshy Radicle 77 © 2014 Pearson Education, Inc. ! The flowers of a given plant species are synchronized to appear at a specific time of the year to promote outbreeding 78 © 2014 Pearson Education, Inc. 79 © 2014 Pearson Education, Inc. 80 © 2014 Pearson Education, Inc. Figure 38.10 Figure 38.11 Figure 38.11a Carpels ! Fruits are classified based on their developmental origin ! Simple fruits develop from a single or several fused carpels Carpels Stamen Petal Style Ovary Stamen Stigma Ovule Pea flower ! Aggregate fruits result from a single flower with multiple separate carpels Raspberry flower Stigma Seed Ovary Pineapple inflorescence Each segment develops from the carpel of one flower Stamen Pea fruit (a) Simple fruit Stamen Sepal Ovule Carpel (fruitlet) ! Multiple fruits develop from a group of flowers called an inflorescence Flowers Pineapple fruit (c) Multiple fruit Pea flower Apple flower Remains of stamens and styles Sepals 81 Raspberry flower Carpel (fruitlet) Stigma Ovary Receptacle Apple fruit (d) Accessory fruit 82 © 2014 Pearson Education, Inc. Ovule Seed Seed Raspberry fruit (b) Aggregate fruit Stigma Ovary (in receptacle) Stamen Pea fruit © 2014 Pearson Education, Inc. Stamen Ovary Stamen Stigma 83 © 2014 Pearson Education, Inc. (a) Simple fruit Raspberry fruit (b) Aggregate fruit 84 © 2014 Pearson Education, Inc. Figure 38.11b Flowers Petal Stigma Style ! Fruit dispersal mechanisms include ! Water Ovary (in receptacle) ! Wind Apple flower Remains of stamens and styles Sepals Seed Pineapple fruit (c) Multiple fruit ! An accessory fruit contains other floral parts in addition to ovaries Stamen Sepal Ovule Pineapple inflorescence Each segment develops from the carpel of one flower Animation: Fruit Development ! Animals Receptacle Apple fruit 85 (d) Accessory fruit © 2014 Pearson Education, Inc. 86 © 2014 Pearson Education, Inc. Figure 38.12a 87 © 2014 Pearson Education, Inc. Figure 38.12aa 88 © 2014 Pearson Education, Inc. Figure 38.12ab Figure 38.12ac Dispersal by water Dispersal by wind Giant seed of the tropical Asian climbing gourd Alsomitra macrocarpa Coconut seed embryo, endosperm, and endocarp inside buoyant husk Dandelion fruit Dandelion fruit Dandelion “seeds” (actually one-seeded fruits) 89 Winged fruit of a maple Giant seed of the tropical Asian climbing gourd Alsomitra macrocarpa Coconut seed embryo, endosperm, and endocarp inside buoyant husk 90 Dandelion “seeds” (actually one-seeded fruits) 91 92 Tumbleweed © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Figure 38.12ad © 2014 Pearson Education, Inc. Figure 38.12ae Figure 38.12b Figure 38.12ba Dispersal by animals Fruit of puncture vine (Tribulus terrestris) Winged fruit of a maple © 2014 Pearson Education, Inc. Squirrel hoarding seeds or fruits underground Fruit of puncture vine (Tribulus terrestris) Tumbleweed Ant carrying seed with attached “food body” 93 94 95 96 Seeds dispersed in black bear feces © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Figure 38.12bb Figure 38.12bc Figure 38.12bd Concept 38.2: Flowering plants reproduce sexually, asexually, or both ! Many angiosperm species reproduce both asexually and sexually ! Sexual reproduction results in offspring that are genetically different from their parents ! Asexual reproduction results in a clone of genetically identical organisms Squirrel hoarding seeds or fruits underground Ant carrying seed with attached “food body” Seeds dispersed in black bear feces 97 © 2014 Pearson Education, Inc. 98 © 2014 Pearson Education, Inc. 99 © 2014 Pearson Education, Inc. 100 © 2014 Pearson Education, Inc. Figure 38.13 Mechanisms of Asexual Reproduction Advantages and Disadvantages of Asexual and Sexual Reproduction ! Fragmentation, separation of a parent plant into parts that develop into whole plants, is a very common type of asexual reproduction ! Apomixis is the asexual production of seeds from a diploid cell ! In some species, a parent plant’s root system gives rise to adventitious shoots that become separate shoot systems ! Asexual reproduction is also called vegetative reproduction because progeny arise from mature vegetative fragments ! All genetic material is passed to the progeny ! Asexual reproduction can be beneficial to a successful plant in a stable environment ! However, a clone of plants is vulnerable to local extinction if there is an environmental change 101 © 2014 Pearson Education, Inc. 102 © 2014 Pearson Education, Inc. 103 © 2014 Pearson Education, Inc. 104 © 2014 Pearson Education, Inc. Figure 38.14 Figure 38.14a Mechanisms That Prevent Self-Fertilization ! Sexual reproduction generates genetic variation that makes evolutionary adaptation possible ! Many angiosperms have mechanisms that make it difficult or impossible for a flower to self-fertilize ! However, only a fraction of seedlings survive ! Dioecious species have staminate and carpellate flowers on separate plants ! Some flowers can self-fertilize to ensure that every ovule will develop into a seed (a) Staminate flowers (left) and carpellate flowers (right) of a dioecious species ! However, many species have evolved mechanisms to prevent selfing Stamens Styles Styles 105 © 2014 Pearson Education, Inc. 106 © 2014 Pearson Education, Inc. Figure 38.14b Thrum flower Staminate flower Stamens Pin flower (b) Thrum and pin flowers 107 © 2014 Pearson Education, Inc. 108 © 2014 Pearson Education, Inc. Figure 38.14c ! Others have stamens and carpels that mature at different times or are arranged to prevent selfing ! The most common is self-incompatibility, a plant’s ability to reject its own pollen ! Researchers are unraveling the molecular mechanisms involved in self-incompatibility Stamens Styles Styles Stamens ! Some plants reject pollen that has an S-gene matching an allele in the stigma cells Thrum flower Carpellate flower 109 © 2014 Pearson Education, Inc. Pin flower ! Recognition of self pollen triggers a signal transduction pathway leading to a block in growth of a pollen tube 110 © 2014 Pearson Education, Inc. 111 © 2014 Pearson Education, Inc. 112 © 2014 Pearson Education, Inc. Totipotency, Vegetative Reproduction, and Tissue Culture Vegetative Propagation and Grafting Test-Tube Cloning and Related Techniques ! Totipotent cells, those that can divide and asexually generate a clone of the original organism, are common in plants ! Vegetative reproduction that is facilitated or induced by humans is called vegetative propagation ! Humans have devised methods for asexual propagation of angiosperms ! Many kinds of plants are asexually reproduced from plant fragments called cuttings ! Most methods are based on the ability of plants to form adventitious roots or shoots ! A callus is a mass of dividing, undifferentiated totipotent cells that forms where a stem is cut and produces adventitious roots 113 © 2014 Pearson Education, Inc. ! A twig or bud can be grafted onto a plant of a closely related species or variety ! Plant biologists have adopted in vitro methods to create and clone novel plant varieties ! The stock provides the root system ! A callus of undifferentiated totipotent cells can sprout shoots and roots in response to plant hormones ! The scion is grafted onto the stock 114 © 2014 Pearson Education, Inc. 115 © 2014 Pearson Education, Inc. 116 © 2014 Pearson Education, Inc. Figure 38.15 Figure 38.16 Concept 38.3: People modify crops by breeding and genetic engineering ! Some pathogenic viruses can be eliminated by excising virus-free apical meristems for tissue culture ! People have intervened in the reproduction and genetic makeup of plants for thousands of years ! Plant tissue culture also facilitates the production of genetically modified (GM) plants (a) (b) (c) ! Hybridization is common in nature and has been used by breeders to introduce new genes ! Maize, a product of artificial selection, is a staple in many developing countries Developing root 117 © 2014 Pearson Education, Inc. 118 © 2014 Pearson Education, Inc. Figure 38.16a 119 120 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Plant Breeding Plant Biotechnology and Genetic Engineering Figure 38.16b ! Mutations can arise spontaneously or can be induced by breeders ! Plant biotechnology has two meanings ! In a general sense, it refers to innovations in the use of plants to make useful products ! Plants with beneficial mutations are used in breeding experiments ! In a specific sense, it refers to use of GM organisms in agriculture and industry ! Desirable traits can be introduced from different species or genera 121 © 2014 Pearson Education, Inc. 122 © 2014 Pearson Education, Inc. ! Transgenic organisms are those that have been engineered to express a gene from another species 123 © 2014 Pearson Education, Inc. 124 © 2014 Pearson Education, Inc. Figure 38.17 Figure 38.18 Reducing World Hunger and Malnutrition ! Genetically modified plants may increase the quality and quantity of food worldwide ! Nutritional quality of plants is being improved ! For example, “Golden Rice” is a transgenic variety being developed to address vitamin A deficiencies among the world’s poor ! Some transgenic crops have been developed to produce the Bt toxin, which is toxic to insect pests ! Other crops are able to tolerate herbicides or resist specific diseases Non-Bt maize 125 © 2014 Pearson Education, Inc. ! For example, transgenic cassava have increased levels of iron and beta-carotene and reduced cyanide-producing chemicals Bt maize 126 © 2014 Pearson Education, Inc. 127 © 2014 Pearson Education, Inc. 128 © 2014 Pearson Education, Inc. Reducing Fossil Fuel Dependency The Debate over Plant Biotechnology ! Biofuels are fuels derived from living biomass, the total mass of organic matter in a group of organisms Issues of Human Health ! Some biologists are concerned about risks of releasing GM organisms (GMOs) into the environment ! One concern is that genetic engineering may transfer allergens from a gene source to a plant used for food ! Biofuels can be produced by rapidly growing crops such as switchgrass and poplar ! Widespread adoption of Bt cotton in India has led to a 41% decrease in insecticide use and an 80% reduction in acute poisoning cases ! Some GMOs have health benefits ! For example, maize that produces the Bt toxin has 90% less of a cancer-causing toxin than non-Bt corn ! Biofuels would reduce the net emission of CO2, a greenhouse gas ! Bt maize has less insect damage and lower infection by Fusarium fungus that produces the cancer-causing toxin 129 130 © 2014 Pearson Education, Inc. © 2014 Pearson Education, Inc. Possible Effects on Nontarget Organisms Addressing the Problem of Transgene Escape 131 © 2014 Pearson Education, Inc. 132 © 2014 Pearson Education, Inc. Figure 38.UN01a ! Many ecologists are concerned that the growing of GM crops might have unforeseen effects on nontarget organisms ! Perhaps the most serious concern is the possibility of introduced genes escaping into related weeds through crop-to-weed hybridization ! Efforts are underway to prevent this by introducing ! Male sterility ! Apomixis ! This could result in “superweeds” that would be resistant to many herbicides ! Transgenes into chloroplast DNA (not transferred by pollen) ! Strict self-pollination 133 © 2014 Pearson Education, Inc. 134 © 2014 Pearson Education, Inc. Figure 38.UN01aa 135 © 2014 Pearson Education, Inc. Figure 38.UN01ab 136 © 2014 Pearson Education, Inc. Figure 38.UN01b Figure 38.UN02 Tube nucleus One sperm will fuse with the egg, forming a zygote (2n). One sperm cell will fuse with the 2 polar nuclei, forming an endosperm nucleus (3n). 137 © 2014 Pearson Education, Inc. Figure 38.UN03 141 © 2014 Pearson Education, Inc. 138 © 2014 Pearson Education, Inc. 139 © 2014 Pearson Education, Inc. 140 © 2014 Pearson Education, Inc.