LECTURE CHAPTER 18 - GYMNOSPERMS

Know the Pine life cycle I. EVOLUTIONARY ADVANCES (Part of this is not in your book) A. EVOLUTION OF THE SEED Gymnosperm means naked seed [Greek for gymnos (naked) and sperma (seed)] This refers to the fact that the ovules in gymnosperms are not encased in layers of parental tissue and do not, therefore, form fruits. You can think of gymnosperms, therefore, as fruitless seed plants. Pollen goes directly to ovules rather than to a stigma (as in flowering plants) The development of seeds and pollen that led to the evolution of gymnosperms represents a major evolutionary advancement . This change involves the evolution of increasingly protected gametophytes and increasing independence from water for sexual reproduction. Only cycads and the maiden hair tree have flagellated sperm at all, and all the gymnosperms have their pollen dispersed by wind, animals, or water. These reproductive changes were accompanied by a number of changes in vegetative structures such as stomata, thicker cuticles, and tougher tissue types that help to minimize water loss, all of which were favored by environmental changes as the earth became cooler and drier. Seed ferns, cycads, and conifers all speciated prolifically, and the Jurassic period, for example, is often called the Age of Cycads. Of these once dominant groups, however, only the conifers are abundant today. The earliest seeds were from the late Devonian about 360 million years ago. Ideas about how seeds arose are based on interpretation of the earliest fossils that look like seeds in the late Devonian, about 40 million years after vascular plants appeared (400 million years ago). Fossils show a megasporangium surrounded by a protective integument-like layer. Fossils from later in the Devonian are more seedlike. The megaspore is progressively more enclosed by an integumentary layer (diagram of paleozoic adaptations towards enclosure of the seed Fig 20-5). The eintugment later becomes the seed coat. Seeds represent two sporophyte generations and the gametophyte generation. All seed plants are heterosporous (2 kinds of spores, a macrospore and a microspore). Integuments are sporophyte tissue = seed coat after fertilization. The megasporangium is sporophyte tissue. A single cell in the megasporangium undergoes meiosis. Three of the resulting megaspores disintegrate leaving a functional megaspore. The functional megaspore undergoes repeated nuclear divisions without cytoplasmic division, then cell walls form around each nucleus. Two-200 archegonia develop, each containing a single egg. The megaspore is gametophyte generation (formed by meiosis) = nucellus. The fertilized egg becomes the next sporophyte generation (Zygote --> embryo). There may be one or more embryos (several embryos in a seed = polyembryony. Seeds are retained on the plant after fertilization and are protective for the next sporophyte generation. Embryos (sporophyte generation) develop in the seeds. Delayed maturation until conditions are right must have had a significant evolutionary advantage. It would be a distinct survival advantage if the new sporophyte could use the photosynthetic and absorptive capacity of the previous sporophyte generation. This was accomplished by retaining the megaspore and allowing the megagametophyte to develop inside the megasporophyte. The simplest seed bearing structures are in ginko and yew. Seeds born singly at the ends of stalks. Conifers produce seeds in complex strobili (Smallest = Juniper, Largest = cycads). The seed producing cones of conifers can be referred to as megastrobili. The pollen producing cones can be referred to as microstrobili.

B. EVOLUTIONARY ADVANCES: POLLEN 1. Pollen grains formed by reduction of the male gametophyte and its more secure retention within a protective spore wall. a. See Selagenella and other seedless vascular plants where the male gametophyte is mostly enclosed in a spore wall. b. Botanists distinguish true pollen from microspores by where they germinate. (1) spores germinate from the tetrad scar (place where spore was attached to other meiotic products) (2) Pollen germinates opposite the scar (3) first true pollen didnt appear until the Mesozoic, 150 million years after the first appearance of seeds 2. The Gymnosperm Pollen Grain 1. Pollen grains evolved through the reduction of the male gametophyte from the ancestral thallus of seedless plants. Evidence for this lies in the fact that there are still remnants of older structures which have not been entirely eliminated and are still part of the male gametophyte after the germination of pollen grains. a. One or two cells that are remnants of the gametophyte thallus (prothallial cells) b. A generative cell divides to form a stalk cell (antheredial stalk) and a body cell (antheridium) c. The body cells forms the sperm nuclei II. PROGYMNOSPERMS The oldest known seeds are from the late Devonian, 360 million years ago. During the next 50 million years, a wide array of seed-bearing plants evolved, including the seed ferns, cordaites and conifers. One group of now extinct plants were the progymnosperms. Progymnosperms are considered to be the likely progenitors of gymnosperms. The progymnosperms had a bifacial vascular cambium (one that produces both secondary xylem and secondary phloem) which is characteristic of seed plants. III. THE ECONOMIC IMPORTANCE OF GYMNOSPERMS (Not in your book) A. Gymnosperms are second only to angiosperms in economic importance B. 75% of worlds timber and much of the pulp for paper C. American use 2000,000 tons of paper each day D. Douglas fir (Pseudotsuga menziesii) is second only to sequoia in size and is prized for wood. The timbering of this tree and sequoia has virtually destroyed old growth forests E. Spruce wood is used for sound board in violins because the trachids give good resonance F. Wood products include resin from which turpentine and rosin are derived (called naval stores) G. The huge kauri pines produce a mixture of resins called dammar which is used in varnishes 1. Dammar is also called amber 2. Fossil amber contains remnants of prehistoric life IV. LIVING GYMNOSPERMS A. In general, gymnosperms are trees or shrubs with woody stems, though some are vine-like. They are not a large group with only about 720 species in 65 genera, but they are highly diverse in the structure of their reproductive parts. Microsporaniate strobili, for example, range from flower-like structures in Edphedra to densely packed cones as in most pines, to loosely packed sporangia as in Gingko. Pollen-producing structures are called microsporangia because they produce microspores, and egg (megaspore) producing structures are called megasporangia. Micro- and megasporangia are born on megaphylls (the scales of the cones). B. All but the Gnetophytes lack vessels. In this respect they resemble the seedless vascular plants. Vessels are tube-like structures in the xylem composed of elongate cells placed end to end and connected by pores. Its function is to conduct water and minerals. Found in nearly all angiosperms and a few other vascular plants)

C. Gymnosperms include 1. Phylum Ginkophyta = Ginko 2. Phylum Cycadophyta = Cycads 3. Phylum Coniferophyta = conifers 4. Phylum Gnetophyta (Ephedra and Gnetum) I. PHYLUM CONIFEROPHYTA A. Informally called conifers to indicate that it bears cones 1. Most numerous of the gymnosperms today 2. 50 genera and 550 species 3. Tallest plant is redwood Sequoia sempervirens of coastal California and Oregon can obtain heights up to 117 meters 4. Also includes pines, firs and spruces 5. Leaves have drought-resistant features which may be related to diversification of the division during the dry cold Permian. B. Genus Pinus 1. Most familiar 2. Dominate wide stretches of North America and Eurasia 3. 90 species of pines 4. Needle-like leaves born in bundles or fascicles which are characteristic of the species. 5. Most pines retain needles for 2-4 years 6. Bristlecone pine may be retained for 45 years and are photosynthetically active during this time C. Pine life cycle 1. Pollination occurs. The pollen grains reach the pollination droplet 2. Pollen grains drawn into the ovule, micropyle closes and seed cones close up 3. Meiosis begins in the megaspore 1 month later 4. It takes about 13 months for the female gametophyte to develop from the megaspore. During this time, the pollen tunes digest most of the megasporangial tissue 5. A month before the female gametophyte is fully cellular, the generative cell in each male gametophyte divides and forms a stalk cell and a generative cell. 6. Several days before the pollen tube reaches the female gametophyte, the body cell splits into two sperm cells. The archegonium forms an egg 7. The seed cone is larger and woodier but it remains closed while embryos develop. The embryo divides to form many nuclei which gradually become cellular. (another 6 months) 8. When mature, the cones gradually become dry, open and release seeds 9. Total cycle is 18 months but can vary from 14 to 24 months D. Genus Aibes (Firs) E. Genus Picea (spruces) F. Genus Tusga (hemlock) H. Genus Pseudotsuga (Douglas fir) I. Cupressus (cypresses) J. Family Taxaceae (Yews) 1. Ovules not born in cones but born in fleshy cup-like structure 2. Contain a toxic substance and are a major cause of poisoning of children but rarely fatal 3. Bright color of the berries attracts birds which ingest and spread the seeds K. Family Taxodiaceae 1. Widespread during the Tertiary 2. Scattered populations remain today 3. Sequoia sempervirens (Coast Redwood) 4. Sequoia giganteum (Big tree) 5. Taxodium (Bald cypress) -Southeastern US and Mexico 6. Metasequoia (Dawn redwood) a. Most abundant conifer in the late cretaceous to the Miocene

b. First described from fossil material by the paleobotanist Shigeru Miki in 1941 c. Three years later, the Chinese forester visited a remote village in Sichuan Province in China and discovered a huge tree of a kind he had never seen before. d. Villagers had built a temple at its base. e. He collected needles and cones. Studies indicated it was a remnant of the once great Metasequoia forest, a living fossil f. Seeds have been collected and planted in parks and gardens throughout the world. VI. PHYLUM CYCADOPHYTA A. Ten genera and 100 species of cycad B. Primarily tropical C. All species are dioecious (separate male and female plants) D. Two species native to the US, both in the genus Zamia (Florida) E. Palm-like leaves unlike other gymnosperms F. Roots have nodules containing nitrogen-fixing cyanobacterium G. Strobili are simple. Seeds like those of Ginkgo H. Also have flagellated sperm I. do not produce cones VII. PHYLUM GINKOPHYTA A. Ginkgo biloba is the only living representative of a very ancient division of plants 1. Virtually unchanged for 80 million years 2. No wild stands 3. Tree was cultivated in temple grounds in China and distributed from there 4. Resistant to pollution and can grow in cities B. Ginkgo is exclusively dioecious (Male bearing plants and female or seed bearing plants) C. Flagellated sperm but flagella not needed for pollination D. Ginkos produce a single large seed in a thick fleshy integument. E. Easily recognized by open branching fan-shaped leaves F. Unlike other gymnosperms, it is deciduous VIII. PHYLUM GNETOPHYTA 1. These organisms are unique among the gymnosperms because it undergoes double fertilization. 2. Three genera and 71 species a. Ephedra (40 species) (1) Makes a tea that is a stimulant (ephedra) b. Gnetum (30 species) (1) tropical vines or trees) c. Welwitschia (1 species) (1) Desert plant with two leaves that continue to grow and split 3. Fertilization is unusual a. Tubular growths from the eggs grow towards the pollen tunes. Fertilization occurs in the united structure b. After fertilization, the second sperm fuses with another cell of the female gametophyte (double fertilization) c. Double fertilization otherwise is found only in the angiosperms