Lecture III.6. Plants. Simplified plant phylogeny. The word “plants” (Kingdom Plantae) is here taken to mean “embryophytes” (aka “land plants”), which may be defined as multicellular photoauto- trophs in which one of the life cycle stages is an embryo. This definition excludes green algae (chlorophytes), which can be multicellular, but which lack such a stage. Introduction. “Plants” = “embryophytes”. 1. Multicellular. 2. Chlorophyll a and b cap- ture energy from light. An even simpler phylogeny. 3. Store carbohydrates. 4. Develop from an embryo pro- tected by parental tissue. Charophyte algae the sister group. 1. Use CaCo3 for support. 2. Multicellular. Differentiated into rhizoids, thalli, branchlets. 3. Male (antheridia), female repro- ductive structures (oogonia). 4. Spores protected from desic- cation by sporopollenin, im- A stonewort. Note portant spore and pollen grain the orange male sex organs (antheridia). coat component. 2 Embryophytes can be sequentially subdivided as follows: 1. All Embryophytes: a. Non-tracheophytes. Vascular (water / nutrient conducting) tissue absent: Liverworts, hornworts, mosses. b. Tracheophytes. Vascular tissue present. 2. Tracheophytes: a. Seeds absent: Clubmosses, ferns and related. b. Seeds present: Gymnosperms, angiosperms and related. 3. Angiosperms (flowering plants): a. Monocots. One seed leaf. Gras- ses, palm trees, etc. b. Dicots. Two seed leaves. The majority of plant species. 3 Above. A more detailed plant phylogeny. Note the principal synapomorphies: Ability to live on land; vascular tissue; seeds and flowers. Right. If one equates “plants” with embryo- phytes, the presence of chloro- phyll a and b is a shared an- cestral character, also called a plesiomorphy. 4 Alternation of Generations – Review. Multicellular haploid and diploid individuals. Multicellular sporophyte (2n) 1. Develops mitotically from a diploid zygote. 2. Produces haploid spores Alternating generations in em- by meiosis. bryophytes. Compare with Lec- ture 3.2, p. 21. 3. Spores contained in struc- tures called sporangia. Multicellular gametophyte (n) 1. Develops mitotically from a haploid spore. 2. Produces haploid gametes – also by mitosis. Fusion of two gametes results in the formation of a diploid zygote from which the sporophyte develops, etc. 5 Alternation of Generations in Plants. In most plants, gametes produced by structures called gametangia. Two kinds: 1. Antheridia produce sperm. 2. Archegonia produce eggs (one per archegonium). Gametangia (singular, gametangium) protect the gametes from desiccation and mechanical damage. The embryo develops in, and is nourished by, the archegonium (plural archegonia). 6 Archegonial Zygote Retention. 1. Archegonium the site of fer- tilization. 2. Functions as a womb, in which sporophyte develops. 3. Specialized transfer cells function as a placenta. Micrograph of a liverwort gametophyte showing two 4. In flowering plants (angio- developing sporophytes and sperms), these functions surrounding transfer cells. performed by other struc- tures. Charophyte algae (stone- worts) have archegonial re- tention of the zygote. 1. But these algae are hap- lontic – only the zygote is diploid. Charophyte life cycle. The zygote is retained in the fe- 2. Therefore, if charophytes male gametangium, which is embryophyte sister group, called an oogonium. multicellular sporophyte a derived trait. 7 Moss and fern life cycles. In both groups, sperm cells swim to the egg. In mosses, the gametophyte dominates; in ferns, the sporophyte. 8 In non-tracheophytes, such as mosses, the gameto- phyte is the “plant.” The sporophyte remains attached to the gametophyte on which it is nutritionally dependent. In tracheophytes, the sporophyte is the “plant”. Embryophyte evolution marked by gametophyte reduc- tion and sporophyte enlargement. Progressive reduction of the gametophyte and enlargement of the sporophyte. a. In hornworts (non-tracheophytes), the sporo- phyte sits atop, and is nourished by, the photosynthetic game- tophyte. b. In horsetails (non-seed tracheophytes), the gameto- phyte (left) is much smaller than the sporophyte (right). 9 Evolutionary Time Line. Cambrian. Molecular phylogenies suggest Cambrian / Precambrian origins. Ordovician. Non-tracheophytes. Silurian. Possible tracheophytes. Devonian. 1. Tracheophytes for sure. 2. Trees / forests. 3. Seed plants. Carboniferous. Conifers. Cretaceous. 1. Angiosperms. 2. Darwin’s “abominable mystery”. Cenozoic. Angiosperms dominate. Grasslands extensive. 10 Above Left. Cooksonia (Silurian). Some species may have had xylem and phloem. Note the terminal sporangia (spore-bearing structures. Living plant was about 1½ inches tall. Above Right. Thursophyton (Devonian) with scale-like leaflets reminiscent of living clubmosses (primitive tracheophytes). Next Page. A Devo- nian forest. Acanthostega, a labyrinthodont amphibian, is shown in the foreground. The proposed (G. J. Retallack, 2011. J. Geol. 119: 235-258) origin of tetrapods in flooded woodlands supports an “eat here” hypothesis as opposed Alfred Romer’s earlier suggestion that land dwelling was a response to increasing arid- ity. Still another scenario (J. A. Clack. 2007. Int. Comp, Biol. 47: 510–523) links tetrapod evolution to increasing global anoxia, which would have been more of a problem in stagnant waters than on land. 11 12 Coevolution with Other Organisms. Fungi – mycorrhizal associations provide nutrients. Herbivores. 1. Defense, e.g., phytolyths. 2. Chemical defenses – “secondary compounds.” Pollinators. 1. Flowers attract pollinators. Root tip, root hairs (circled arrows) and mycorrhizae. 2. Examples: a. Bee flowers – yellow, strong scent, concentrat- ed nectar. b. Hummingbird flowers – red, tubular c. Hawkmoth flowers – long, white tubes. Seed dispersers (animals) – e.g., ingestion / elimination. Datura and hawkmoth. Mixed interactions – Hawk- moth adults pollinate Datura; larvae eat the leaves. 13 Yucca–Yucca Moth. An Obligate Mutualism. Fertilization requires services of the moth. 1. Pollen agglutinated1 into masses called pollinia. 2. Ovary superior, i.e., above the pollinia. Female moth. 1. Collects pollinia with ten- Yucca moth gathering pollen. tacles that only she has. 2. Chews them up to free the pollen grains 3. Pushes masticated pollinia down stigmatic canal. 4. Injects eggs into the plant’s ovary with special- ized ovipositor. 5. Go here to watch 1. Developing larvae eat the immature seeds; exit thru fruit wall and pupate in the soil near base of plant. Top. Yucca moth on flower. 2. Yucca plants selectively Bottom. a. seed pod; b. seed abort fruits with too many pod with larval exit holes; c. developing larvae. larva inside seed row. 1 Pollen grains stuck together. 14 Transition to Land. Embryophyte ancestors lived in fresh water – facilitated fertilization by flagellated sperm. Terrestrial existence offered abundant resources: 1. Light. 2. CO2. And it presented problems: 1. Desiccation / gas exchange. 2. Support / transport. 3. Acquiring water / nutrients from the soil. 4. Reproduction – getting the sperm to the egg. 15 Desiccation / Gas Exchange. Waxy cuticle – prevents evaporative water loss. Stomata are pores that permit gas exchange. 1. Each pore surrounded by a pair of guard cells. 2. When the guard cells swell, the pore opens. a. Normally, pores open during the day; close at night. b. When the plant is water-stressed, stomata remain closed during the day. Transverse section of a leaf. Cuticle (upper and lower surface) prevents water loss. Stomata (lower surface) permit gas ex- change via guard cells that swell when exposed to light if the leaf is adequately hydrated. 16 Stomatal opening and closing. In response to light and water avail- ability, potassium ions, 푲+, are pumped into the guard cells. Water follows. This causes the guard cells to bow outward, thereby open- ing the pore. The structural basis of this response is two-fold: 1. Radially oriented cellulose microfibrils prevent guard cell di- ameter from increasing. As a result, the response to water entry is stretching. 2. The thinner outer guard cell walls stretch more easily than the thicker inner walls. This causes the cells bow away from each as shown in the figure. 17 Transport and Support. Vascular plants have specialized structures to transport water, minerals and sugars throughout the plant. 1. Xylem conducts water / minerals from roots to leaves. a. Series of continuous “pipes” formed from dead cells. b. Developed gradually in the course of plant evolution. 2. Xylem also provides support via secondary cell wall. Evolution of xylem in embryophytes. Secondary cell wall formed of a complex biopolymer called lignin, provides strength. Tracheids (cells with pores in the secondary cell wall) and vessel elements (cells with larger openings penetrating both primary and secondary cell walls) enhance transport. Phylogenetic note: Vessel elements evolved inde- pendently in gymnosperms and angiosperms. 18 Water Transport / Support in Non-Tracheophytes. 1. Some, e.g., mosses, have simple conducting cells that lack the perforated secondary cell walls of vascu- lar plant tracheids. 2. Many a. Grow in dense moist masses through which water moves by capillary action. b. Have leaf-like structures that catch and hold water. c. Small enough for nutrients to circulate by diffusion. 3. No support, they just sprawl. 19 Water Transport in Tracheophytes. 1. In aquatic environment, plant surrounded by water. a. Cytosol ~ isotonic with seawater. b. Cell wall prevents membrane rupture