1. LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson © 2011 Pearson Education, Inc. Lectures by Erin Barley Kathleen Fitzpatrick Angiosperm Reproduction and Biotechnology Chapter 38

2. Overview: Flowers of Deceit Insects help angiosperms to reproduce sexually with distant members of their own species –For example, male Campsoscolia wasps mistake Ophrys flowers for females and attempt to mate with them –The flower is pollinated in the process –Unusually, the flower does not produce nectar and the male receives no benefit Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings © 2011 Pearson Education, Inc.

3. Figure 38.1

4. Many angiosperms lure insects with nectar; both plant and pollinator benefit Mutualistic symbioses are common between plants and other species Angiosperms can reproduce sexually and asexually Angiosperms are the most important group of plants in terrestrial ecosystems and in agriculture Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings © 2011 Pearson Education, Inc.

5. Concept 38.1: Flowers, double fertilization, and fruits are unique features of the angiosperm life cycle Plant lifecycles are characterized by the alternation between a multicellular haploid (n) generation and a multicellular diploid (2n) generation Diploid sporophytes (2n) produce spores (n) by meiosis; these grow into haploid gametophytes (n) Gametophytes produce haploid gametes (n) by mitosis; fertilization of gametes produces a sporophyte © 2011 Pearson Education, Inc.

6. Video: Flower Blooming (time lapse)

7. In angiosperms, the sporophyte is the dominant generation, the large plant that we see The gametophytes are reduced in size and depend on the sporophyte for nutrients The angiosperm life cycle is characterized by “three Fs”: flowers, double fertilization, and fruits © 2011 Pearson Education, Inc.

8. Video: Flower Plant Life Cycle (time lapse)

9. Figure 38.2 Stamen Anther Filament Petal Receptacle Stigma Style Ovary Carpel Sepal (a) Structure of an idealized flower Simplified angiosperm life cycle (b) Key Haploid (n) Diploid (2n) Anther Pollen tube Germinated pollen grain (n) (male gametophyte) Ovary Ovule Embryo sac (n) (female gametophyte) Egg (n) Sperm (n) FERTILIZATION Zygote (2n) Mature sporophyte plant (2n) Germinating seed Seed Simple fruit Embryo (2n) (sporophyte)

10. Figure 38.2a Stamen Anther Filament Petal Receptacle Stigma Style Ovary Carpel Sepal (a) Structure of an idealized flower

11. Figure 38.2b Simplified angiosperm life cycle (b) Key Haploid (n) Diploid (2n) Anther Pollen tube Germinated pollen grain (n) (male gametophyte) Ovary Ovule Embryo sac (n) (female gametophyte) Egg (n) Sperm (n) FERTILIZATION Zygote (2n) Mature sporophyte plant (2n) Germinating seed Seed Simple fruit Embryo (2n) (sporophyte)

12. Flower Structure and Function Flowers are the reproductive shoots of the angiosperm sporophyte; they attach to a part of the stem called the receptacle Flowers consist of four floral organs: sepals, petals, stamens, and carpels Stamens and carpels are reproductive organs; sepals and petals are sterile © 2011 Pearson Education, Inc.

13. A stamen consists of a filament topped by an anther with pollen sacs that produce pollen A carpel has a long style with a stigma on which pollen may land At the base of the style is an ovary containing one or more ovules A single carpel or group of fused carpels is called a pistil © 2011 Pearson Education, Inc.

14. Complete flowers contain all four floral organs Incomplete flowers lack one or more floral organs, for example stamens or carpels Clusters of flowers are called inflorescences © 2011 Pearson Education, Inc.

15. Development of Male Gametophytes in Pollen Grains Pollen develops from microspores within the microsporangia, or pollen sacs, of anthers Each microspore undergoes mitosis to produce two cells: the generative cell and the tube cell A pollen grain consists of the two-celled male gametophyte and the spore wall © 2011 Pearson Education, Inc.

16. If pollination succeeds, a pollen grain produces a pollen tube that grows down into the ovary and discharges two sperm cells near the embryo sac © 2011 Pearson Education, Inc.

17. Video: Bee Pollinating

18. © 2011 Pearson Education, Inc. Video: Bat Pollinating Agave Plant

19. Development of a male gametophyte (in pollen grain) (a) Microsporangium (pollen sac) (b)Development of a female gametophyte (embryo sac) Microsporocyte Microspores (4) Each of 4 microspores Generative cell (will form 2 sperm) (LM) 75  m 20  m 100  m MEIOSIS MITOSIS Male gametophyte (in pollen grain) Nucleus of tube cell Ragweed pollen grain (colorized SEM) Key to labels Haploid (n) Diploid (2n) (LM) Embryo sac Ovule Megasporangium Megasporocyte Integuments Micropyle Surviving megaspore Antipodal cells (3) Polar nuclei (2) Egg (1) Synergids (2) Ovule Integuments Female gametophyte (embryo sac) Figure 38.3

20. Development of a male gametophyte (in pollen grain) (a) Microsporangium (pollen sac) Microsporocyte Microspores (4) Each of 4 microspores Generative cell (will form 2 sperm) (LM) 75  m 20  m MEIOSIS MITOSIS Male gametophyte (in pollen grain) Nucleus of tube cell Ragweed pollen grain (colorized SEM) Key to labels Haploid (n) Diploid (2n) Figure 38.3a

21. Development of Female Gametophytes (Embryo Sacs) The embryo sac, or female gametophyte, develops within the ovule Within an ovule, two integuments surround a megasporangium One cell in the megasporangium undergoes meiosis, producing four megaspores, only one of which survives The megaspore divides, producing a large cell with eight nuclei © 2011 Pearson Education, Inc.

22. This cell is partitioned into a multicellular female gametophyte, the embryo sac © 2011 Pearson Education, Inc.

23. Figure 38.3b (b) Development of a female gametophyte (embryo sac) 100  m MEIOSIS MITOSIS Key to labels Haploid (n) Diploid (2n) (LM) Embryo sac Ovule Megasporangium Megasporocyte Integuments Micropyle Surviving megaspore Antipodal cells (3) Polar nuclei (2) Egg (1) Synergids (2) Ovule Integuments Female gametophyte (embryo sac)

24. Figure 38.3c Generative cell (will form 2 sperm) 75  m Nucleus of tube cell (LM)

25. Figure 38.3d 20  m Ragweed pollen grain (colorized SEM)

26. Figure 38.3e 100  m (LM) Embryo sac

27. Pollination In angiosperms, pollination is the transfer of pollen from an anther to a stigma Pollination can be by wind, water, or animals Wind-pollinated species (e.g., grasses and many trees) release large amounts of pollen © 2011 Pearson Education, Inc.

28. Abiotic Pollination by Wind Pollination by Bees Hazel staminate flowers (stamens only) Hazel carpellate flower (carpels only) Common dandelion under normal light Common dandelion under ultraviolet light Figure 38.4a

29. Hazel staminate flowers (stamens only) Figure 38.4aa

30. Figure 38.4ab Hazel carpellate flower (carpels only)

31. Figure 38.4ac Common dandelion under normal light

32. Figure 38.4ad Common dandelion under ultraviolet light

33. Pollination by Moths and Butterflies Blowfly on carrion flower Pollination by Flies Pollination by Bats Moth on yucca flower Long-nosed bat feeding on cactus flower at night Hummingbird drinking nectar of columbine flower Pollination by Birds Stigma Anther Moth Fly egg Figure 38.4b

34. Figure 38.4ba Moth on yucca flower Stigma Anther Moth

35. Figure 38.4bb Blowfly on carrion flower Fly egg

36. Figure 38.4bc Long-nosed bat feeding on cactus flower at night

37. Figure 38.4bd Hummingbird drinking nectar of columbine flower

38. Coevolution of Flower and Pollinator Coevolution is the evolution of interacting species in response to changes in each other Many flowering plants have coevolved with specific pollinators The shapes and sizes of flowers often correspond to the pollen transporting parts of their animal pollinators –For example, Darwin correctly predicted a moth with a 28 cm long tongue based on the morphology of a particular flower © 2011 Pearson Education, Inc.

39. Figure 38.5

40. Double Fertilization After landing on a receptive stigma, a pollen grain produces a pollen tube that extends between the cells of the style toward the ovary Double fertilization results from the discharge of two sperm from the pollen tube into the embryo sac One sperm fertilizes the egg, and the other combines with the polar nuclei, giving rise to the triploid food-storing Plant Fertilization endosperm (3n) © 2011 Pearson Education, Inc.

41. Animation: Plant Fertilization Right-click slide / select “Play”

42. Figure 38.6-1 Stigma Pollen tube 1 2 sperm Style Ovary Ovule Micropyle Pollen grain Polar nuclei Egg

43. Figure 38.6-2 Stigma Pollen tube 21 2 sperm Style Ovary Ovule Micropyle Pollen grain Polar nuclei Egg Ovule Polar nuclei Egg Synergid 2 sperm

44. Figure 38.6-3 Stigma Pollen tube 231 2 sperm Style Ovary Ovule Micropyle Pollen grain Polar nuclei Egg Ovule Polar nuclei Egg Synergid 2 sperm Endosperm nucleus (3n) (2 polar nuclei plus sperm) Zygote (2n)

45. Seed Development, Form, and Function After double fertilization, each ovule develops into a seed The ovary develops into a fruit enclosing the seed(s) © 2011 Pearson Education, Inc.

46. Endosperm Development Endosperm development usually precedes embryo development In most monocots and some eudicots, endosperm stores nutrients that can be used by the seedling In other eudicots, the food reserves of the endosperm are exported to the cotyledons © 2011 Pearson Education, Inc.

47. Embryo Development The first mitotic division of the zygote splits the fertilized egg into a basal cell and a terminal cell The basal cell produces a multicellular suspensor, which anchors the embryo to the parent plant The terminal cell gives rise to most of the embryo The cotyledons form and the embryo elongates © 2011 Pearson Education, Inc.

48. Animation: Seed Development Right-click slide / select “Play”

49. Figure 38.7 Ovule Endosperm nucleus Integuments Zygote Terminal cell Basal cell Proembryo Suspensor Basal cell Cotyledons Shoot apex Root apex Suspensor Seed coat Endosperm

50. Ovule Endosperm nucleus Integuments Zygote Terminal cell Basal cell Figure 38.7a

51. Figure 38.7b Proembryo Suspensor Basal cell Cotyledons Shoot apex Root apex Suspensor Seed coat Endosperm

52. Structure of the Mature Seed The embryo and its food supply are enclosed by a hard, protective seed coat The seed enters a state of dormancy A mature seed is only about 5–15% water © 2011 Pearson Education, Inc.

53. In some eudicots, such as the common garden bean, the embryo consists of the embryonic axis attached to two thick cotyledons (seed leaves) 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 The plumule comprises the epicotyl, young leaves, and shoot apical meristem © 2011 Pearson Education, Inc.

54. Figure 38.8 Seed coat Radicle Epicotyl Hypocotyl Cotyledons (a) Common garden bean, a eudicot with thick cotyledons (b) Castor bean, a eudicot with thin cotyledons (c) Maize, a monocot Seed coat Endosperm Cotyledons Epicotyl Hypocotyl Radicle Hypocotyl Epicotyl Endosperm Pericarp fused with seed coat Scutellum (cotyledon) Coleoptile Coleorhiza

55. Figure 38.8a Seed coat Radicle Epicotyl Hypocotyl Cotyledons (a) Common garden bean, a eudicot with thick cotyledons

56. The seeds of some eudicots, such as castor beans, have thin cotyledons © 2011 Pearson Education, Inc.

57. Figure 38.8b (b) Castor bean, a eudicot with thin cotyledons Seed coat Endosperm Cotyledons Epicotyl Hypocotyl Radicle

58. 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 © 2011 Pearson Education, Inc.

59. Figure 38.8c (c) Maize, a monocot Radicle Hypocotyl Epicotyl Endosperm Pericarp fused with seed coat Scutellum (cotyledon) Coleoptile Coleorhiza

60. Seed Dormancy: An Adaptation for Tough Times 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 © 2011 Pearson Education, Inc.

61. Seed Germination and Seedling Development Germination depends on imbibition, the uptake of water due to low water potential of the dry seed The radicle (embryonic root) emerges first Next, the shoot tip breaks through the soil surface © 2011 Pearson Education, Inc.

62. In many eudicots, a hook forms in the hypocotyl, and growth pushes the hook above ground Light causes the hook to straighten and pull the cotyledons and shoot tip up © 2011 Pearson Education, Inc.

63. Figure 38.9 Foliage leaves Cotyledon Hypocotyl Cotyledon Hypocotyl Radicle Seed coat Epicotyl Cotyledon Hypocotyl (a) Common garden bean Foliage leaves Coleoptile Radicle (b) Maize

64. Figure 38.9a Foliage leaves Cotyledon Hypocotyl Cotyledon Hypocotyl Radicle Seed coat Epicotyl Cotyledon Hypocotyl (a) Common garden bean

65. In maize and other grasses, which are monocots, the coleoptile pushes up through the soil © 2011 Pearson Education, Inc.

66. Figure 38.9b Foliage leaves Coleoptile Radicle (b) Maize

67. Fruit Form and Function A fruit develops from the ovary It protects the enclosed seeds and aids in seed dispersal by wind or animals A fruit may be classified as dry, if the ovary dries out at maturity, or fleshy, if the ovary becomes thick, soft, and sweet at maturity © 2011 Pearson Education, Inc.

68. Animation: Fruit Development Right-click slide / select “Play”

69. Fruits are also classified by their development –Simple, a single or several fused carpels –Aggregate, a single flower with multiple separate carpels –Multiple, a group of flowers called an inflorescence © 2011 Pearson Education, Inc.

70. Figure 38.10 Stamen Ovary Stigma Ovule Pea flower Seed Pea fruit (a) Simple fruit (b) Aggregate fruit (c) Multiple fruit (d) Accessory fruit Carpels Stamen Raspberry flower Carpel (fruitlet) Stigma Ovary Stamen Raspberry fruit Flower Pineapple inflorescence Each segment develops from the carpel of one flower Pineapple fruit Stigma Petal Style Stamen Sepal Ovule Ovary (in receptacle) Apple flower Remains of stamens and styles Sepals Seed Receptacle Apple fruit

71. Figure 38.10a Stamen Ovary Stigma Ovule Pea flower Seed Pea fruit (a) Simple fruit (b) Aggregate fruit Carpels Stamen Raspberry flower Carpel (fruitlet) Stigma Ovary Stamen Raspberry fruit

72. Figure 38.10b (c) Multiple fruit (d) Accessory fruit Flower Pineapple inflorescence Each segment develops from the carpel of one flower Pineapple fruit Stigma Petal Style Stamen Sepal Ovule Ovary (in receptacle) Apple flower Remains of stamens and styles Sepals Seed Receptacle Apple fruit

73. An accessory fruit contains other floral parts in addition to ovaries © 2011 Pearson Education, Inc.

74. Fruit dispersal mechanisms include –Water –Wind –Animals © 2011 Pearson Education, Inc.

75. Dispersal by Wind Dandelion “seeds” (actually one-seeded fruits) Winged fruit of a maple Dandelion fruit Tumbleweed Dispersal by Water Winged seed of the tropical Asian climbing gourd Alsomitra macrocarpa Coconut seed embryo, endosperm, and endocarp inside buoyant husk Figure 38.11a

76. Figure 38.11aa Coconut seed embryo, endosperm, and endocarp inside buoyant husk

77. Figure 38.11ab Winged seed of the tropical Asian climbing gourd Alsomitra macrocarpa

78. Figure 38.11ac Dandelion “seeds” (actually one-seeded fruits) Dandelion fruit

79. Figure 38.11ad Winged fruit of a maple

80. Figure 38.11ae Tumbleweed

81. Figure 38.11b Dispersal by Animals Fruit of puncture vine (Tribulus terrestris) Squirrel hoarding seeds or fruits underground Ant carrying seed with nutritious “food body” to its nest Seeds dispersed in black bear feces

82. Figure 38.11ba Fruit of puncture vine (Tribulus terrestris)

83. Figure 38.11bb Squirrel hoarding seeds or fruits underground

84. Figure 38.11bc Seeds dispersed in black bear feces

85. Figure 38.11bd Ant carrying seed with nutritious “food body” to its nest

86. 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 © 2011 Pearson Education, Inc.

87. Mechanisms of Asexual Reproduction Fragmentation, separation of a parent plant into parts that develop into whole plants, is a very common type of asexual reproduction In some species, a parent plant’s root system gives rise to adventitious shoots that become separate shoot systems © 2011 Pearson Education, Inc.

88. Figure 38.12

89. Apomixis is the asexual production of seeds from a diploid cell © 2011 Pearson Education, Inc.

90. Advantages and Disadvantages of Asexual Versus Sexual Reproduction Asexual reproduction is also called vegetative reproduction 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 © 2011 Pearson Education, Inc.

91. Sexual reproduction generates genetic variation that makes evolutionary adaptation possible However, only a fraction of seedlings survive Some flowers can self-fertilize to ensure that every ovule will develop into a seed Many species have evolved mechanisms to prevent selfing © 2011 Pearson Education, Inc.

92. Mechanisms That Prevent Self-Fertilization Many angiosperms have mechanisms that make it difficult or impossible for a flower to self-fertilize Dioecious species have staminate and carpellate flowers on separate plants © 2011 Pearson Education, Inc.

93. Figure 38.13 Staminate flowers (left) and carpellate flowers (right) of a dioecious species (a) (b) Thrum and pin flowers Thrum flower Pin flower Stamens Styles

94. Figure 38.13a Staminate flowers

95. Others have stamens and carpels that mature at different times or are arranged to prevent selfing © 2011 Pearson Education, Inc.

96. Figure 38.13b Carpellate flowers

97. Figure 38.13c Thrum flowerPin flower Stamens Styles

98. 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 Some plants reject pollen that has an S-gene matching an allele in the stigma cells Recognition of self pollen triggers a signal transduction pathway leading to a block in growth of a pollen tube © 2011 Pearson Education, Inc.

99. Vegetative Propagation and Agriculture Humans have devised methods for asexual propagation of angiosperms Most methods are based on the ability of plants to form adventitious roots or shoots © 2011 Pearson Education, Inc.

100. Clones from Cuttings Many kinds of plants are asexually reproduced from plant fragments called cuttings A callus is a mass of dividing undifferentiated cells that forms where a stem is cut and produces adventitious roots Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings © 2011 Pearson Education, Inc.

101. Grafting A twig or bud can be grafted onto a plant of a closely related species or variety The stock provides the root system The scion is grafted onto the stock © 2011 Pearson Education, Inc.

102. Test-Tube Cloning and Related Techniques Plant biologists have adopted in vitro methods to create and clone novel plant varieties A callus of undifferentiated cells can sprout shoots and roots in response to plant hormones © 2011 Pearson Education, Inc.

103. Figure 38.14 Developing root (a) (b) (c)

104. Transgenic plants are genetically modified (GM) to express a gene from another organism Protoplast fusion is used to create hybrid plants by fusing protoplasts, plant cells with their cell walls removed © 2011 Pearson Education, Inc.

105. Figure 38.15 50  m

106. Concept 38.3: Humans modify crops by breeding and genetic engineering Humans have intervened in the reproduction and genetic makeup of plants for thousands of years 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 © 2011 Pearson Education, Inc.

107. Figure 38.16

108. Figure 38.16a

109. Figure 38.16b

110. Plant Breeding Mutations can arise spontaneously or can be induced by breeders Plants with beneficial mutations are used in breeding experiments Desirable traits can be introduced from different species or genera The grain triticale is derived from a successful cross between wheat and rye © 2011 Pearson Education, Inc.

111. Plant Biotechnology and Genetic Engineering Plant biotechnology has two meanings –In a general sense, it refers to innovations in the use of plants to make useful products –In a specific sense, it refers to use of GM organisms in agriculture and industry Modern plant biotechnology is not limited to transfer of genes between closely related species or varieties of the same species © 2011 Pearson Education, Inc.

112. Reducing World Hunger and Malnutrition Genetically modified plants may increase the quality and quantity of food worldwide Transgenic crops have been developed that –Produce proteins to defend them against insect pests –Tolerate herbicides –Resist specific diseases © 2011 Pearson Education, Inc.

113. 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 © 2011 Pearson Education, Inc.

114. Figure 38.17 Cassava roots harvested in Thailand

115. Biofuels are made by the fermentation and distillation of plant materials such as cellulose Biofuels can be produced by rapidly growing crops such as switchgrass and poplar Biofuels would reduce the net emission of CO 2, a greenhouse gas The environmental implications of biofuels are controversial Reducing Fossil Fuel Dependency © 2011 Pearson Education, Inc.

116. The Debate over Plant Biotechnology Some biologists are concerned about risks of releasing GM organisms (GMOs) into the environment © 2011 Pearson Education, Inc.

117. Issues of Human Health One concern is that genetic engineering may transfer allergens from a gene source to a plant used for food 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 –Bt maize has less insect damage and lower infection by Fusarium fungus that produces the cancer-causing toxin © 2011 Pearson Education, Inc.

118. GMO opponents advocate for clear labeling of all GMO foods © 2011 Pearson Education, Inc.

119. Possible Effects on Nontarget Organisms Many ecologists are concerned that the growing of GM crops might have unforeseen effects on nontarget organisms © 2011 Pearson Education, Inc.

120. Addressing the Problem of Transgene Escape Perhaps the most serious concern is the possibility of introduced genes escaping into related weeds through crop-to-weed hybridization This could result in “superweeds” that would be resistant to many herbicides © 2011 Pearson Education, Inc.

121. Efforts are underway to prevent this by introducing –Male sterility –Apomixis –Transgenes into chloroplast DNA (not transferred by pollen) –Strict self-pollination © 2011 Pearson Education, Inc.

122. Figure 38.UN01 Endosperm nucleus (3n) (2 polar nuclei plus sperm) Zygote (2n) (egg plus sperm)

123. Figure 38.UN02