Section 1: Pollination enables gametes to come together within a flower

1. Section 1: Pollination enables gametes to come together within a flower Chapter 38: Angiosperm Reproduction and Biotechnology

2. Review: The Angiosperm Life Cycle Recall that: -The life cycles of plants have an alteration of generationsbetween haploid and diploid states. -In angiosperms, the sporophyteis the dominate state. -Pollination is achieved by wind, water, or animals.

3. Flower Structure -Flowers are the reproductive shoots of the angiosperm sporophyte. -They are typically composed of four whorls of highly modified leaves called floral organs, which are separated by short internodes. -Flowers are determinate shoots, meaning they cease growing after the flower and fruit are formed

4. The Floral Organs • Consist of the sepals, petals, stamens, and carpels that are attached to a part of the stem called the receptacle. • Reproductive organs:stamensand carpels • Sterile organs: sepals and petals • Sepals : a modified leaf that closes and protects the floral bud before it opens • Petals : modified leaf of a flowering plant; often colorful parts of a flower that advertise it to insects and other pollinators

5. Subparts of Floral Organs • Stamen : consists of a stalk called a filament and a terminal structure called the anther • anther : contains chambers called pollen sacs in which pollen is produced • Carpel : consists of an ovary at its base, and a long slender neck called the style • style: has a sticky structure called a stigma at the top that serves as a platform for pollen to land • ovary: contains one or more ovules • pistil : refers to a group of fused carpels

6. Floral Variations • Complete flowers have all the basic floral organs. • Incomplete flowers lack one or more of these floral organs. SymmetryOvary Location Floral Distribution • Flowers can differ in symmetry. • They can have bilateral symmetry, in which they can be divided into two equal parts by an imaginary line. • They can also have radical symmetry, in which any imaginary line through the central axis divides the flower into two equal parts • The location of a flower’s ovary may vary in relation to the stamens, petals, and sepals. • An ovary is called superior if those parts are attached below it; semi-inferior if they are attached alongside it; and inferior if they are attached above it. • Floral distribution can also differ. • Some species have individual flowers, while others have clusters called inflorescences.

7. Floral Variations (cont’d) Reproductive Variations • -In most incomplete flowers, stamens and carpels are wither absent or nonfunctional. • Incomplete flowers that have only function stamen are called staminate, and those with only functional carpels are called carpellate. • If staminate and carpellate flowers are on the same species, the species is said to be monoecious. • -A dioecious species has staminate flowers and carpellate flowers on separate plants.

8. Terms: Gametophyte Development and Pollination • Microspores : spores from a heterosporous plant species that develops into a male gametophyte • Megaspores : spore from a heterosporous plant species that develops into a female gametophyte

9. Gametophyte Development and Pollination (cont’d) Development of a male gametophyte (pollen grain). Pollen sacs develop within the microsporangia (pollen sacs) of anthers at the tips of the stamens. • Each of the microsporangia contains diploid microsporocytes (microspore mother cells). • Each microsporocyte divides by meiosis, producing four haploid microspores, each of which develops into a pollen grain. • A pollen grain becomes a mature male gametophyte when it generative nucleus divides and forms two sperm. This usually occurs after a pollen gain lands on the stigma of a carpel and the pollen tube begins to grow.

10. Gametophyte Development and Pollination (cont’d) Development of a female gametophyte(embryo sac). The embryo sac develops within an ovule, itself enclosed by the ovary at the base of a carpel. • Within the ovule’s megasporangium is a large diploid cell called the megasporocyte. • The megasporocyte divides by meiosis and gives rise to four haploid cells, but in most species only one of these survives as the megaspore. • 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).

11. Gametophyte Development and Pollination (cont’d) • Pollination, the transfer of pollen from anther to stigma, is the first step in a chain of events that can lead to fertilization. • Plants can rely on: air, water, or animals for pollination.

12. Mechanisms That Prevent Self-Fertilization • Many angiosperms have mechanisms that make it difficult or impossible for self-fertilization or “selfing” to occur. • This contributes to genetic variety by ensuring that the sperm and eggs come from different parents. • The most common anti-selfing mechanism is self-incompatibility, the ability of a plant to reject its own pollen and sometimes the pollen of a closely related relative.

13. Mechanisms That Prevent Self-Fertilization (cont’d) • Recognition of “self” pollen is based on genes for self-incompatibility, called S genes. • If a pollen grain has an allele that matches an allele of the stigma on which it lands, self-recognition blocks growth by either : gametophytic self-compatibility or sporophytic self-compatibility. • Gametophytic self-compatibility: The S –allele in the pollen genome governs the blocking of fertilization. • Ex.) S1 pollen grain from an S1S2 parental sporophyte will fail to fertilize eggs of an S1S2 flower but will fertilize an S2S3 flower. An S2 pollen grain would not fertilize either flower • Involves enzymatic destruction of RNA within a rudimentary pollen tube

14. Mechanisms That Prevent Self-Fertilization (cont’d) • Sporophytic self-compatibility: fertilization is blocked by S –allele gene products in tissues of the parental sporophyte that adhere to the pollen wall. • Ex.) neither an S1 nor S2pollen grain from an S1S2 parental sporophyte will fertilize eggs of an S1S2 flower or S2S3 flower • Involves a signal transduction pathway in epidermal cells of the stigma

15. 38.2After fertilization, ovules develop into seeds and ovaries into fruits. A look at fertilization and its products: seeds and fruits.

16. Double Fertilization • After landing on a receptive stigma, a pollen grain absorbs moisture and germinates • Produces a pollen tube that extends down between the cells of the style towards the ovary • Nucleus of generative cell divides by mitosis and forms two sperm • Tip of pollen tube enters ovary, discharges two sperm near embryo sac

17. Double Fertilization • Angiosperm life cycle: • One sperm fertilizes egg and forms zygote • Other sperm combines with two polar nuclei to form triploid nucleus • Triploid nucleus in large central cell, cell gives rise to endosperm (food-storage) • Two sperm cells joined with different nuclei of the embryo sac is double fertilization • Prevents angiosperms from squandering nutrients

18. Double Fertilization • First cellular event after gamete fusion is increase in cytoplasmic calcium levels of the egg • Also occurs during animal gamete fusion • Establishment of a block to polyspermy (fertilization of an egg by more than one sperm cell) also similar to animals

19. From Ovule to Seed • After double fertilization each ovule becomes a seed and the ovary becomes a fruit enclosing the seed • Seeds stockpile proteins, oils, and starch and become sugar sinks • Storage function of endosperm is eventually taken over by swelling cotyledons of embryo

20. Endosperm Development • Precedes embryo development • After double fertilization, triploid nucleus divides forming multinucleate supercell • This liquid mass (endosperm) becomes multicellular when cytokinesis forms membranes between nuclei • Naked cells produce cell walls and endosperm becomes solid • Ex: coconut milk vs. coconut meat

21. Embryo Development • First mitotic division of the zygote is transverse, splitting the fertilized egg into a basal cell and a terminal cell • Terminal cell gives rise to most of embryo • Basal cell continues to divide transversely producing thread of cells (suspensor) which anchors embryo to parent • Suspensor functions in transfer of nutrients and in protection • Terminal cell divides several times, forms proembryo attached to suspensor • Cotyledon begins to form as bumps on proembryo

22. Structure of the Mature Seed • Seed dehydrates in final stages, making water content 5-15% of weight • Embryo becomes dormant • Embryo and food supply are enclosed by seed coat • Hypocotyl (embryonic axis) terminates in the radicle (embryonic root) • Portion above where cotyledons attach is the epicotyl • Embryo of a monocot has a single cotyledon • Grass family has scutellum (specialized cotyledon) which is thin with large surface area for absorption • Embryo of grass seed enclosed by two sheaths • Coleoptile covers young shoot, coleorhiza covers young root

23. From Ovary to Fruit • Fruit protects enclosed seeds and aids in their dispersal • Fertilization triggers hormonal changes that cause ovary-fruit transformation • Ovary wall becomes pericarp (thickened fruit wall) • Most fruits, derived from a single carpel or several fused carpels, are simple fruits • Ex: peach, pea pod, nut • Aggregate fruit results from single flower with more than one separate carpel; clustered fruitlets • Ex: raspberry • Multiple fruit develops from inflorescence, group of flowers clustered tightly together; walls thicken and fuse • Ex: pineapple

24. From Ovary to Fruit • Sometimes other floral parts contribute to fruit (accessory fruits) • Apple flowers: fruit embedded in receptacle and the fleshy part is derived mainly from receptacle; only core comes from ovary • Strawberries: aggregate fruit consisting of enlarged receptacle embedded with tiny one-seeded fruits • Fruit ripens when seeds complete development • Ripening of dry fruit involves aging & drying out of tissues • Complex hormone interactions enable ripening of fleshy fruits

25. Seed Germination • As seed matures, it dehydrates and enters dormancy • Extremely low metabolic rate • Suspension of growth and development • Some seeds germinate as soon as they enter suitable environment • Others remain dormant until receiving specific environmental cue

26. Seed Dormancy • Seed dormancy is for increased chance of germination at advantageous time and place • Breaking dormancy requires specific conditions • Ex: desert plants may germinate after substantial rainfall, and not mild drizzle • Many seeds require intense heat to break dormancy, or extended exposure to cold • Some require light and some have coats that must be weakened by chemical attack (in digestion) • Soil has bank of ungerminated seeds that accumulate and reappear after disaster

27. From Seed to Seedling • Germination of seeds depends on imbibition (uptake of water due to the low water potential of dry seed) • Imbibing water causes seed to expand and rupture coat; triggers metabolic changes in embryo to enable resumed growth • First organ to emerge from germinating seed is radicle (embryonic root) followed by shoot tip • Followed by hypocotyl, cotyledons, epicotyl

28. From Seed to Seedling • Monocots (maize and other grasses) use different method • Coleoptile (sheath enclosing shoot) pushes upward followed by shoot tip through tubular coleoptile and out coleoptile’s tip • In germination, tough seed gives way to fragile seedling exposed to harsh conditions • Only a fraction of seedlings survive in the wild • Production of seeds in enormous numbers compensates for this

29. Many flowering plants clone themselves by asexual reproduction

30. Asexual Reproduction Exact copies are derived from one parent Has no genetic recombination Many plants produce sexually and asexually Ex: seedless, gymnosperms and some angiosperms Vegetative reproduction- asexual plants are more mature then when sexually reproduced Beneficial when in an unstable environment

31. Mechanisms of Asexual Reproduction Asexual reproduction is an extension of the capacity for indeterminate growth Plants have undifferentiated cells which can divide, sustaining or renewing growth indefinitely Detached vegetative fragments can develop into whole offspring Fragmentation-separation of plant parts which grow into individual entire plants Common form of asexual reproduction

32. Mechanisms of Asexual Reproduction Seeds produced without fertilization or pollination is a process called apomixis Only one parent, no sperm and egg Diploid cell in ovule matures into seeds-clones of parent Seeds are dispersed, although seed dispersal us typically a sexual process

33. Vegetative Propagation and Agriculture Genetic modification  occurs by incorporating foreign genes to get better characteristics Clones from cuttings  asexual reproduction using plant fragments After a cut at end of shoot or stem mass of undifferentiated cells called callus forms Cells differentiate and can become whole plant Grafting-modification that is a result of cuttings combines best qualities of plants in vivo stock: root system scion: twig grafted onto stock

34. Vegetative Propagation and Agriculture Plants can be created or cloned in vitro Cells cultured and then begin to divide so placed in soil to grow into a plant Whole plants can grow from culturing small pieces of parent tissue Genetic modification also occurs in vitro Transgenic-organisms engineered to express a gene from another species

35. Vegetative Propagation and Agriculture Protoplast fusion Protoplasts-plant cells with cell walls removed Screened for mutations looking to improve the agricultural value of the plant Different species protoplasts can be fused Otherwise reproductively incompatible Now are “hybrids” ex- potato and wild potato combined to acquire resistance to a weed killer

36. Plant Biotechnology is Transforming Agriculture!!! “Biotechnology has the potential to bring tremendous benefits …” -John Prescott

37. Artificial Selection • Humans have developed selective plant traits since at least 10,000 years ago in the Neolithic Stone Age • Maize • Regular maize is poor source of protein • Low tryptophan and lysine, 2/8 essential amino acids • Mutant maize opaque-2 have much higher levels of tryptophan and lysine • Problems: opaque-2 maize kernels have soft endosperm  1) harder to harvest 2) more prone to attack by pests

38. Artificial Selection • Maize, cont. • Using hybridization and artificial selection, scientists created a “super maize” with a hard endosperm and sufficient protein nutrition • Using modern genetic engineering techniques, scientists can transfer genes from completely unrelated species

39. Reducing World Hunger and Malnutrition • 40,000 people die a day from malnutrition • Some feel it is because the poor cannot afford food • Others say the world is overpopulated, and it has already reached its carrying capacity • The main goal of biotechnology is to increase crop production • Methods: • 1- Transgenic Crops with Toxins • Ex.) hybrid cotton, maize, and potatoes • Contain the gene Bt toxin from bacterium Bacillus thuringiensis • Bt toxin : • 1- controls the number of serious insect pests • 2- is harmless to vertebrates • 3- reduces the need for insecticide spraying

40. Reducing World Hunger and Malnutrition • 2- Resistant Transgenic Crops • Ex.) transgenic soybeans, sugar cane, and wheat • Resistant to herbicides and “weed killers” • Ex.) Ring-spot virus resistant papaya plants were introduced in Hawaii • 3- Increased Nutritional Plant Quality • Ex.) “golden” rice • Has a few daffodil genes • Increases vitamin A, helping prevent blindness

41. The Debate Over Plant Biotechnology • Issues of Human Health • Many people are concerned that genetic engineering may pass allergens from gene source to plant • In fact, there is no evidence that genetically modified organisms (GMOs) have adverse effects on human health usually positive effects! • Ex.) Bt Maize has less cancer-causing agents, fumonisin • Those against GMOs want: • 1- labeling of GMO products • 2- Separate transport, processing, and storage of GMO products

42. The Debate Over Plant Biotechnology • Possible Effects on Non-target Organisms • STUDY: Caterpillars of monarch butterflies died after consumption of milkweed leaves heavily dusted with pollen from Bt maize • It was eventually found that other plant parts with Bt toxin caused death • Despite the negative effects of Bt maize, the alternative to insect control would be insecticide spraying, having an even greater impact on non-target organisms

43. The Debate Over Plant Biotechnology • Addressing the Problem of Transgene Escape • transgene escape – the escape of genes from transgenic crops into related weeds via crop-to-weed hybridization • The major fear of biotechnologists is that a “super weed” will develop • Because of transgene escape, scientists are trying to: • 1- breed male sterility into transgenic crops no viable pollen would be produced • 2- engineer transgene into chloroplast DNA of crop • Would work because chloroplast DNA is often inherited strictly from maternal plants, so transgenes in chloroplasts cannot be transferred by pollen

44. The Debate Over Plant Biotechnology • “Terminator technology” is another possible solution for transgene escape • GMOs that undergo the terminator process grow normally until last stages of seed or pollen maturation • Then, “terminator” protein, toxic to plants, is activated in nearly mature seed or pollen, making the seeds or pollen unviable • These proteins are produced only if original seeds are pretreated with a specific chemical