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(5) Chapter 38 Angiosperm Reproduction and Biotechnology 被子植物生殖與生物技術 PowerPoint Lectures for Biology, Seventh Edition Neil Campbell and Jane Reece Lectures by Chris Romero Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings To Be or Not to Be (??????) • Overview: To Seed or Not to Seed • The parasitic plant Rafflesia arnoldii – Produces enormous flowers that can produce up to 4 million seeds Figure 38.1 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Key Concepts Concept 38.1: Pollination enables gametes to come together within a flower (受粉作用使得花 中的配子聚在一起) Concept 38.2: After fertilization, ovules develop into seeds and ovaries into fruits Concept 38.3: Many flowering plants clone (複製) themselves by asexual reproduction Concept 38.4: Plant biotechnology is transforming agriculture Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 38.1: Pollination enables gametes to come together within a flower (受粉作用使 得花中的配子聚在一起) • In angiosperms, the dominant sporophyte (孢 子體) – Produces spores that develop within flowers into male gametophytes (pollen grains) (雄配子體、花粉粒) – Produces female gametophytes (embryo sacs) (雌配子體、胚囊) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Flower Structure • Flowers – Are the reproductive shoots of the angiosperm sporophyte (被子植物配子體) – Are composed of four floral organs: • Sepals (萼片) 雄蕊Stamen • Petals (花瓣) 花藥 Anther Filament 花絲 • Stamens (雄蕊) • Carpels (雌蕊) Carpel雌蕊 Ovary卵房 Sepal萼片 花瓣Petal Receptacle 花托 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 柱頭 Stigma Style 花柱 被子植物花的多樣性 • Many variations in floral structure have evolved during the 140 million years of angiosperm history SYMMETRY OVARY LOCATION Bilateral symmetry (orchid) Radial symmetry (daffodil) FLORAL DISTRIBUTION Superior ovary Sepal Lupine inflorescence Semi-inferior Inferior ovary ovary Fused petals Sunflower inflorescence REPRODUCTIVE VARIATIONS Maize, a monoecious species Figure 38.3 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Dioecious Sagittaria latifolia (common arrowhead) Gametophyte Development and Pollination • In angiosperms (被子植物) – Pollination (受粉作用) is the transfer of pollen from an anther (花藥) to a stigma (柱 頭) – If pollination is successful, a pollen grain (花粉粒) produces a structure called a pollen tube (花粉管), which grows down into the ovary (卵房) and discharges sperm near the embryo sac (胚囊) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • An overview of angiosperm reproduction Stamen Anther Carpel Stigma Style Filament Germinated pollen grain (n) (male gametophyte) on stigma of carpel Anther at tip of stamen Ovary Ovary (base of carpel) Pollen tube Ovule (卵胞) Embryo sac (n) (female gametophyte) Sepal Egg (n) Petal FERTILIZATION Receptacle Sperm (n) (a) An idealized flower. Key Mature sporophyte plant (2n) with flowers Seed Seed (develops from ovule) Zygote (2n) Haploid (n) Diploid (2n) (b) Simplified angiosperm life cycle. See Figure 30.10 for a more detailed version of the life cycle, including meiosis. Figure 38.2a, b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Embryo (2n) (sporophyte) Germinating seed Simple fruit (develops from ovary) • Pollen (花粉)---Develops from microspores (雄 孢子) within the sporangia (孢子囊) of anthers (a) Development of a male gametophyte (pollen grain)雄配子體(花粉粒)的發育 Pollen sac (microsporangium) (1) Each one of the microsporangia contains diploid microsporocytes (2n) (microspore mother cells). (2) Each microsporocyte divides by meiosis to produce four haploid microspores (n), each of which develops into a pollen grain. (3) A pollen grain becomes a mature male gametophyte when its generative nucleus divides and forms two sperm. This usually occurs after a pollen grain lands on the stigma of a carpel and the pollen tube begins to grow. (See Figure 38.2b.) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings MicroSporocyte MEIOSIS減數分裂 Microspores (4) Each of 4 microspores Generative cell (will form 2 sperm) MITOSIS有絲分裂 Male Gametophyte (pollen grain) Nucleus of tube cell 20 m KEY to labels Haploid (2n) Diploid (2n) 75 m Ragweed pollen grain Figure 38.4a • Embryo sacs---Develop from megaspores within ovules (b) Development of a female gametophyte (embryo sac)雌配子體(胚囊)的發育 Ovule 減數分裂MEIOSIS Megasporangium Megasporocyte Integuments Micropyle Surviving megaspore Female gametophyte (embryo sac) 有絲分裂MITOSIS Ovule Antipodel Cells (3) Polar Nuclei (2) Egg (1) Integuments Synergids (2) Haploid (2n) Diploid (2n) 100 m Key to labels Embryo sac (1) Within the ovule’s megasporangium is a large diploid cell called the megasporocyte (megaspore mother cell). (2) The megasporocyte divides by meiosis and gives rise to four haploid cells, but in most species only one of these survives as the megaspore. (3) Three mitotic divisions of the megaspore form the embryo sac, a multicellular female gametophyte. The ovule now consists of the embryo sac along with the surrounding integuments (protective tissue). Figure 38.4b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mechanisms That Prevent Self-Fertilization (自我受精) • Many angiosperms (被子植物) – Have mechanisms that make it difficult or impossible for a flower to fertilize itself (自我受精) Stigma 柱頭 Stigma 柱頭 Anther with Pollen 有花 粉的花藥 Pin flower Figure 38.5 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Thrum flower • The most common anti-selfing mechanism in flowering plants – Is known as self-incompatibility (自我排斥、自我不 相容), the ability of a plant to reject its own pollen • Researchers are unraveling the molecular mechanisms that are involved in self-incompatibility • Some plants – Reject pollen that has an S-gene matching an allele in the stigma cells • Recognition of self pollen (自家花粉) – Triggers a signal transduction pathway leading to a block (阻礙) in growth of a pollen tube Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 38.2: After fertilization, ovules develop into seeds and ovaries into fruits Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Double Fertilization (雙重受精) • After landing on a receptive stigma (花柱) – A pollen grain (花粉粒) germinates and produces a pollen tube (花粉管) that extends down between the cells of the style () toward the ovary () • The pollen tube (花粉管) – Then discharges two sperm into the embryo sac • In double fertilization (雙重受精) – One sperm fertilizes the egg – The other sperm combines with the polar nuclei (極 核), giving rise to the food-storing endosperm (胚乳) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Growth of the pollen tube and double fertilization (花粉粒)Pollen grain (1) If a pollen grain germinates, a pollen tube grows down the style toward the ovary. (極核)Polar nuclei Egg Stigma (柱頭) Pollen tube (花粉管) 2 sperm Style Ovary Ovule (containing female gametophyte, or embryo sac) Micropyle (2) The pollen tube discharges two sperm into the female gametophyte (embryo sac) within an ovule. (3) One sperm fertilizes the egg, forming the zygote. The other sperm combines with the two polar nuclei of the embryo sac’s large central cell, forming a triploid cell that develops into the nutritive tissue called endosperm. Ovule Polar nuclei Egg Two sperm about to be discharged Endosperm nucleus (3n) (2 polar nuclei plus sperm) Zygote (2n) (egg plus sperm) Figure 38.6 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Development of Ovule and Endosperm • From Ovule to Seed, after double fertilization – Each ovule develops into a seed – The ovary develops into a fruit enclosing the seed(s) • Endosperm Development, usually precedes embryo development – In most monocots and some eudicots, the endosperm stores nutrients that can be used by the seedling after germination • In other eudicots, the food reserves of the endosperm are completely exported to the cotyledons Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Embryo Development • The first mitotic division of the zygote is transverse – Splitting the fertilized egg into a basal cell and a terminal cell Ovule Endosperm nucleus Integuments Zygote Terminal cell Basal cell Zygote Proembryo Suspensor Basal cell Cotyledons Shoot apex Root apex Seed coat Endosperm Figure 38.7 Suspensor Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Structure of the Mature Seed • The embryo and its food supply – Are enclosed by a hard, protective seed coat • In a common garden bean, a eudicot – The embryo consists of the hypocotyl, radicle, and thick cotyledons 種皮 Seed coat Epicotyl 上胚軸 Hypocotyl 下胚軸 Radicle 胚根 Figure 38.8a Cotyledons 子葉 (a) Common garden bean, a eudicot with thick cotyledons. The fleshy cotyledons store food absorbed from the endosperm before the seed germinates. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The seeds of other eudicots, such as castor beans – Have similar structures, but thin cotyledons Seed coat Seed coat Endosperm Endosperm Cotyledons Cotyledons Epicotyl Epicotyl Hypocotyl Hypocotyl Radicle Radicle 種皮 胚乳 子葉 上胚軸 下胚軸 胚根 (b) Castor bean, a eudicot with thin cotyledons. The narrow, membranous cotyledons (shown in edge and flat views) absorb food from the endosperm when the seed germinates. Figure 38.8b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The embryo of a monocot (單子葉) has a single cotyledon, a coleoptile, and a coleorhiza (子葉)Scutellum (cotyledon) (芽鞘) Coleoptile (根鞘) Coleorhiza Pericarp fused with seed coat (種皮) Endosperm (胚乳) Epicotyl (上胚軸) Hypocotyl (下胚軸) Radicle (胚根) (c) Maize, a monocot. Like all monocots, maize has only one cotyledon. Maize and other grasses have a large cotyledon called a scutellum. The rudimentary shoot is sheathed in a structure called the coleoptile, and the coleorhiza covers the young root. Figure 38.8c Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings From Ovary to Fruit • A fruit – Develops from the ovary – Protects the enclosed seeds – Aids in the dispersal of seeds by wind or animals Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Fruits are classified into several types – Depending on their developmental origin Carpels Ovary Stamen Flower Stigma Stamen Ovule Pea flower Raspberry flower Carpel (fruitlet) Seed Stigma Ovary Stamen Pineapple inflorescence Each segment develops from the carpel of one flower Pea fruit Raspberry fruit (a) Simple fruit. A simple fruit develops from a single carpel (or several fused carpels) of one flower (examples: pea, lemon, peanut). (b) Aggregate fruit. An aggregate fruit develops from many separate carpels Figure 38.9a–c of one flower (examples: raspberry, blackberry, strawberry). Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Pineapple fruit (c) Multiple fruit. A multiple fruit develops from many carpels of many flowers (examples: pineapple, fig). Seed Germination (種子萌芽) • As a seed matures – It dehydrates (脫水) and enters a phase referred to as dormancy (休眠) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Seed Dormancy: 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 cues • From Seed to Seedling (從種子到幼苗) – Germination of seeds depends on the physical process called imbibition (浸潤) – The uptake of water due to low water potential of the dry seed Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • The radicle (胚軸) is the first organ to emerge from the germinating seed • In many eudicots, a hook forms in the hypocotyl (下胚軸), and growth pushes the hook above ground Foliage leaves (初生葉) (上胚軸) Hypocotyl (上胚軸) Hypocotyl Cotyledon Cotyledon Epicotyl (上胚軸) Cotyledon (子葉) Hypocotyl (上胚軸) Radicle (胚根) Figure 38.10a Seed coat (種皮) (a)Common garden bean. In common garden beans, straightening of a hook in the hypocotyl pulls the cotyledons from the soil. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Monocots (單子葉植物) – Use a different method for breaking ground when they germinate • The coleoptile (芽鞘) – Pushes upward through the soil and into the air Foliage leaves Coleoptile Coleoptile Radicle Figure 38.10b (b) Maize. In maize and other grasses, the shoot grows straight up through the tube of the coleoptile. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Concept 38.3: Many flowering plants clone (複 製) themselves by asexual reproduction • Many angiosperm species reproduce both asexually and sexually (無性及有性) • Sexual reproduction – Generates the genetic variation that makes evolutionary adaptation possible • Asexual reproduction in plants – Is called vegetative reproduction (營養繁殖) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Mechanisms of Asexual Reproduction (無性生殖的機制) • Fragmentation (裂片) – Is the separation of a parent plant into parts that develop into whole plants – Is one of the most common modes of asexual reproduction • In some species, the root system of a single parent gives rise to many adventitious shoots that become separate shoot systems Figure 38.11 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Vegetative Propagation (營養繁殖) and Agriculture • Humans have devised various methods for asexual propagation of angiosperms • Clones from cuttings (切枝、切條) – Many kinds of plants are asexually reproduced from plant fragments called cuttings • Grafting (架接、接枝) – In a modification of vegetative reproduction from cuttings – A twig or bud from one plant can be grafted onto a plant of a closely related species or a different variety of the same species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Test-Tube Cloning (試管複製) and Related Techniques • Plant biologists have adopted in vitro methods to create and clone novel plant varieties. (a) Just a few parenchyma cells from a carrot gave rise to this callus, a mass of undifferentiated cells. (b) The callus differentiates into an entire plant, with leaves, stems, and roots. Figure 38.12a, b Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • In a process called protoplast fusion (原生 質體融合) – Researchers fuse protoplasts, plant cells with their cell walls removed, to create hybrid plants Vacuole Chloroplast Figure 38.13 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 50 m • Concept 38.4: Plant biotechnology is transforming agriculture • Plant biotechnology has two meanings: – It refers to innovations in the use of plants to make products of use to humans – It refers to the use of genetically modified organisms (GMO, 基因改造生物) in agriculture and industry Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Artificial Selection (人工選擇; 人擇) • Humans have intervened – In the reproduction and genetic makeup of plants for thousands of years • Maize (玉米) – Is a product of artificial selection by humans – Is a staple (xxxxx) in many developing countries, but is a poor source of protein • Interspecific hybridization of plants (植物的種間雜交) – Is common in nature and has been used by breeders, ancient and modern, to introduce new genes Figure 38.14 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Reducing World Hunger and Malnutrition (營養不良) • Genetically modified (GM) plants – Have the potential of increasing the quality and quantity of food worldwide Genetically modified rice GM papaya Figure 38.15 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Ordinary rice Figure 38.16 The Debate over Plant Biotechnology (植物生物技術的爭議) • There are some biologists, particularly ecologists – Who are concerned about the unknown risks associated with the release of GM organisms (GMOs) into the environment • Issues of Human Health – One concern is that genetic engineering may transfer allergens from a gene source to a plant used for food • Possible Effects on Nontarget Organisms – Many ecologists are concerned that the growing of GM crops might have unforeseen effects on nontarget organisms Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings The Debate over Plant Biotechnology (植物生物技術的爭議) • Addressing the Problem of Transgene Escape – Perhaps the most serious concern that some scientists raise about GM crops is the possibility of the introduced genes escaping from a transgenic crop into related weeds through crop-to-weed hybridization (作物雜草雜交) • Despite all the issues associated with GM crops – The benefits should be considered Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 報告完畢 敬請指教 !? !? !? !? !? !? Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings