BIOL 412 — Lecture (Unit 4)
Bryophytes
- 290,000+ known species of land plants: mosses, ferns, gymnosperms, flowering plants
Evolution of Land Plants
- Timeline
- 1.2 bya — thin coating of cyanobacteria and protists, likely close to water
- 500 mya — small plants, fungi, and animals on land
- 470 mya — numerous fungal spore fossils
- 385 mya — taller plants (lignin)
- Plantae clade/supergroup, Viridiplantae, Streptophyta
- Land plants: Embryophytes
- Exclusive of charophytes
- Proteins that synthesize cellulose are arranged in a rosette, with sub-units derived from inside the cell center vs. linear proteins
- Chara — stonewort (calcium carbonate)
- Coleochaete — sporopollenin
- Land plants have various opportunities and challenges
- More sunlight, more CO2 available
- Nutrients no longer bathe plants (mycorrhizae), issues competing for sunlight
- All plants adapt to dry land via
- Alteration of Generations lifecycle
- Oogamous
- vs. charophytes
- spores are protected against dessication and can opt to grow into gametophyte
- Multicellular sporangia that produce “Walled spores”
- Division/differentiation for walls
- Sporopollenin: Spores and pollen greens, tough and resistant, hard to digest
- Multicellular gametangia in most plant phyla
- Multiple cells protecting the egg and sperm being made
- Consistent structure
- Egg producing gametangium is the archegonium
- Flask/pear shaped
- Has a larger venter (egg) and a neck that sperm swims through
- Sperm producing gametangium is the antheridium
- Surrounded by a layer: sterile jacket; both outside on the gametangiumm and on each individual
- Multicellular young sporophyte is retained and nourished by the parent: embryo
- Advancement over single zygotes not protected/nourished
- 3-dimensional apical meristems
- three-dimensional grrowth vs. intertwined filaments (parenchymatous) growth
Bryophytes
- Non-vascular plants; no vascular tissue
- Paraphyletic group/Grade of evolution
- No lignin for conducting cells of xylem → no true vascular tissue
- May have rhizoids: unicellular or filamentous structures that anchor the plant in place; no need for absorptive structure, can dessicate
- No true leaves; no veins/vascular tissue (no roots, stems)
- Gametophyte is the dominant generation
- Bisexual or Unisexual
- Male gametophyte is a antheridium sperm, female is an archegonium sperm
- Releases and swims through a film of water; fertilizes egg and forms a diploid zygote
- Zygote divides through mitosis to form embryo, then sporophyte
- One spore producing structure: sporangium
- Meiosis of diploid cells: sporocytes
- Swimming sperm
- Dominant gametophyte, reduced and dependent sporophyte
Bryophyta
- Mosses phylum
- Leafy area at bottom is dominant gametophyte
- Stalk is the sporophyte; non-dominant, ephemeral
- Three classes of mosses
- Wet/moist areas, cold, volcanic area
- Spiral arrangement of “leaves” around the central axis
- Upright growth supported by thicker cell walls, not lignin
- Can completely dry out and rehydrate; poikilohydry
- Mostly haploid; two part haploid stage
- Haploid spores germinate and form filamentous green mass: protonema(ta)
- Bulbles/buds: main part of gametophyte developing
- Gives rise to leafy gametophyte
- Many are bisexual, but have different stems that are unisexual
- Some are single-sex
- Sexual structures are on top of gametophyte, on top of stem: antheridial/archegonial head
- Sterile hairs
- Archegonial head has longer, stalked structures beneath venter
- Venter is torn in two, then carried upwards and enlarges: calyptra
- Venter falling off allows spore dispersal; leaves cap of sporangium (operculum) and peristome teeth that disperse spores
- Three parts of gametophyte
- Capsule: Sporangium
- Seta: Stalk
- Foot: Anchors to gametophyte
- Characteristics of sporophyte
- Dependent on gametophyte for nutrition; turns brown quickly aand no longer photosynthetic
- True stomata with one cell vs. guard cells; some contrrol over dessication
- Larger sporophytes have specialized cells for water and transport; water hydroids and leptoids
True mosses
- Polytrichum (hair/cat moss)
Peat mosses
- Sphagnum spp.
- Wetlands
- Marginally different gametophyte: “stemmed”/branched/tufted gametophytes
- Leaf layer has two types of cells
- Living cells: green, narrow, networked
- Dead cells: large cells with ridges of thick cell wall and pores
- Holds a lot of water
- Soil mixtures, soil bags
- Contains decay-resistant phenolics
- Antiseptic and anti-microbial properties
- Wound dressings in WW2 (pus, growth of bacteria), Native diapers, soils
- Acidifies habitat
- Prevents decomposition
- Secretes acids, protons
- Layers build up and don’t decompose: carbon remnants; source of fuel
- Floating mat bogs: layer of moss accumulates soil, allowing vegetation floating on open water
- Bogmen left behind, preserved with hair and skin; “Tollund Man”
- Ecological importance: 3% of land mass and accumulate 30% of soil carbon
- vs. dessication via climate change; increased rates of decomposition and carbon release
- Peat: compressed, partially decomposed plant matter
- Peat can buildup in waterlogged soils without moss; more with peat moss
Liverworts
- Hepatophyta spp. — Liver phylum
- Lobed
- “Doctrine of signatures”
- Leafy liverworts and thallose liverworts
- Tropical leafy liverworts: leaflike structures that are tiny; more common
- Thallose liverworts have a thallus instead of large differentiation
- Thallose branches dichotomously/diverges
- Flat on the ground
- ~2cm wide
- Thallus serves as the dominant gametophyte
- Non-green cells on bottom hold water; green cells on top
- No true stomata; only open pores that close when the plant dessicates
- Thallose liverworts have asexual and sexual reproduction
- Gemmae cups/splash cups produced by mitosis: multicellular packets
- Splash cups; grow into new gametophyte thallus
- Reproductive structures are unisexual; can only be discerned after “fruiting”
- Long stalks; antheridial heads/antheridial fours, archegonia heads/fours
- Flat antheridial heads, leafy archegonia heads
- Archegonia heads have hanging archegonia; sperm must be splashed or swum up to fertilize egg
- Generates microscopic sporophyte with a foot, short seta, and large sporangium capsule
- Dispersed spores from top when dry
- Marchantia
Hornworts
- Anthocerophyta spp.
- The horn is the sporophyte; sporophyte is the sporangium
- Short stalk and a short foot
- Sporangium continuously grows, and splits topdown as spores release
- Tetrad: four spores
- True stomata
Seedless vascular plants
- Grade of evolution
- Ancestral vascular tissue; no extant seeds
- Divided into Lycophytes (club mosses, spike mosses, quillworts) and monilophytes (ferns, horsetails, whisk ferns)
- Has lignin in the tracheids and vessel elements in the xylem
- Reconstructed ancestor may not have had vascular tissue (Aglaophyton) but had branched sporophyte stems that could produce multiple sporangia
- Sporophyte is the dominant generation
- Spores aid in dry adaptations: protects pollen
- Various characteristics
- True vascular tissue and lignin in the xylem conducting elements
- Cuticles and true stomata protect water loss and gas exchange
- Sporophyte is the dominant generation and is longer lasting or larger
- Both generations are nutritionally and physically independent
- Gametophytes can be either underground and dependent upon mycorrhizae or aboveground and green
- Sperm are flagellated and need free water for fertilization
- Most plants have roots; some do not
- True stems with vascular tissue (stele)
- True leaves
- Plants have spores and sporangia
- Stems have developing stele: vascular tissue present in the stem or root
- Early development: protostele
- Irregular development
- Solid core of central vascular tissue
- Siphonostele
- A pith with a ring of vascular tissue
- Siphon-like
- May have leaf gaps or no leaf gaps
- Eustele/True stele
- Vascular bundles
- Bundles are able to merge and come apart
- Eustele and siphonostele evolved independent of one another
- Most groups have true leaves with vascular tissue and veins; two types
- Microphyll
- Monovein leaves
- Particular evolutionary line: the enation theory
- a protostele stem without leaves developed flaps of green photosynthetic tissue
- enations: flaps of green photosynthetic tissue that increased surface area for photosynthesis but were not venated
- vascular supply was redirected to enation, developing a vein and a microphyll
- Selaginella kraussiana
- Megaphyll
- Polyvein leaves
- Hymenophyllum tunbridgense
- Telome theory
- Telome: end branches of the megaphyll
- A green stem that dichotomously branched in three-dimensions (bushy) developed equal dichotomous branches
- Overtopping: a main stem developed, with one branch becoming unequally larger while the side branch remaining continued to branch
- Planation: the side branch flattened onto a plane, forming the basis for the veins of the megaphyll leaf
- Webbing: green tissue that was not venated branches out, forming the megaphyll
- Two types of spores/sporangia: homosporous and heterosporous
- Homosporous
- Monodic spore that grows into a bisexual gametophyte
- Gametophyte contains both sexes of gametangia
- Heterosporous
- Two types of spores
- Spores develop into unisexual gametophytes
- Reduced and independent gametophyte, dominant sporophyte
Lycophyta
- Monophyletic phyla
- Lycophytes
- Three extant families
- Phylum was prevalent in Carboniferous era in swamps
- carbon today comes from living plants
- tree-sized lycophytes in swamps and free standing water; present species are very small
- Protosteles and microphylls
- Regardless of actual size of microphylls
- Bean-shaped sporangia
Spike Mosses
- Selaginella moellendorffii; only Selaginella genus
- Selaginellaceae
- Underground rhizomes with true roots
- Aboveground aerial stems with spiral microphylls
- Heterosporous with reduced gametophyte size
- Some spike mosses are tender and flattened (ironlike/planated); some are spiked
- Resurrection plant: xerophytic, can dry completely and rehydrate
- Megasporangium are on megasporophyll and develop megaspores
- Megaspore develop into female gametophytes (megagametophyte) that develop eggs
- Microsporangium are on microsporophyll and develop microspores
- Microspore develop into male gametophytes (microgametophyte) that develop sperm
- Particular lifecycle
- Megaspore develops a sporophyte
- Sporophyte matures devloping roots and strobili
- Strobili contain
- one megasporangium that contains one diploid megasporocyte
- Megasporocyte goes through meiosis and develops four megaspores
- many microsporangium
- many microsporocytes developing four microspores in meiosis
- No particular arrangement off megasporangium and microsporangium
- Free-sporing release
- Microspore dividees by mitosis and eventually develops motile sperm
- Megaspore wall develops a protruding megagametophyte wwith archegonia to fertilize
Club Mosses
- Diphasiastrum tristachyum, Lycopodium spp., Huperzia
- Lycopodiaceae family
- Underground rhizomes with true roots branching
- Aboveground aerial stems with spiral microphylls
- Homosporous and potentially bisexual
- Commonly identified by “clubs”: strobili clusters of sporophylls
- Sporophyte contains particular protostele regardless of sample
- Solid mass of vascular tissue
- Interconnected plates of xylem with phloem borders
- Lateral bean-shaped sporangia
- Sporophylls: leaves that also hold sporangia
- Strobilus: compact arrangement of sporophylls and sporangia
- Sporocytes
- Free-sporing lifecycle: spores are released before gametophytes are generated
Quillworts
- Isoetes gunnii; Isoetes genus, Isoetaceae family
- Wetlands
- Contractile roots that anchor into drying mud
- CAM photosynthesis
Monilophyta
- Monilophytes
- Three subphyla/families
Ferns
- Largest family; ~11,000
- Athyrium filix-femina
- Many species are tropical
- Many species are epiphytic; lives on other plants
- Some epiphytic species are not tropical
- Commensalist
- Siphonosteles: Cylindrical piths that may have leaf gaps
- “Leaves” are fronds; fronds are megaphylls
- Polyvenation
- Fronds are divided into leaflets (pinnae) and can have secondary subdivision; high division
- Fronds are typically the only part above ground; rhizome is belowground and roots branch from rhizomes
- Ferns make “fiddleheads” when young leaves are growing
- Tight scroll of leaf cells
- Differential cell enlargement
- Uncurling and straightening leaves flatten out into mature leaves
- Protects young tissue
- May store toxins
- Numerous “typical” ferns
Filicales
- Majority of ferns in this order
- Leptosporangium unique to Filicales
- Annulus specialization: a row of cells specialized to open up sporangia and catapult spores out
- Homosporous
- Group of sporangia: sorus (sori)
- May be protected by a flap of tissue: indusium
- “Upside-down umbrella” shape
- Indusium shrivels and allows spores to free
- Variety of indusia shapes: bean-shape etc.
- Mature gametophyte is small, heart-shaped
- Base of sporophyte; anchored by rhizoids
- Non-dominant generation
- Fully absorptive tissue
- Notch and tip at center with an archegonia close to the notch and antheridia close to the tip
- Typically bisexual
- Flimsy green one-layer cells
- Sporophyte has notable vascular tissue
Eusporangiate
- Has a eusporangium
- Thick sporangia with no annulus
- Botrychium, Ophioglossum
- large sporangia
- Many tree ferns are eusoporangiate
Water Ferns
- Aquatic
- Heterosporous
- Marsilea, Azolla
Horsetails
- Equisetum telmateia
- “Scouring rushes”
- Equisetum genus
- Some are branched; horse-tail appearance (branches, not leaves)
- Some do not have branches; different stems
- Diverse vegetative and fertile stems
- Fertile stems have strobili
- Large airspaces
- Carinal canals
- Wet areas that require oxygen exchange and water transport between roots and shoots
- Tree-sized horsetails in carboniferous era (360mya)
- Microphylls reduced from megaphylls; different origin of microphylls
- Hollow stems with carinal canals
- Large amounts of silica in epidermis; rough ridges
- Solid jointed nodes; jointed stems that are photosynthetic
- Sporangia are arranged in umbrella-shaped strobili with sporangiophores
- Modified stems: sporangiophores
- Multiple sporangia hang from end of sporangiophores
- Special spore dispersal using elaters
- Spores have ribbonlike parts in spore wall that detach: elaters
- Opens and wing like when dry; wraps around spore when moist
Whiskferns
- Psilotum nudum
- Only Psilotum and Tmesipteris genera
- Formerly “primitive”; enations are hypothesized reduced from true leaves
- Tropical
Seed Plants
- Gymnosperms and angiosperms
- Reduced and dependent gametophyte, dominant sporophyte
- Heterosporous
- No free-sporing behavior before gametophyte development; release after gametophyte development
- Reduction and loss of structures
- Pollen grains and ovules
- Pollen grains transfer in pollination and fertilize; sperm do not have to swim
- Pollination: transfer of pollen from pollen grain to ovary
- Fertilization: sperm fuses with egg
- Ovules become seeds after fertilization; better dispersal units to spores
- Present in Carboniferous era, but greatly diversified during Permian period duee to changes in climate
- Warm/wet to dry/cold change
- Colder, drier, seeds and pollen grains were advantageous
- True secondary growth
- Increased competition for sunlight
Pollen Grains
- Male gametophytes with coating material
- Do not require water for fertilization
- Not free-sporing
- Microspores divide by mitosis to make pollen grains, then release from microsporangia
- Sporopollenin wall; prevents dessication
- Four cells in cone; tetradic
- Transferred by air pollination
- Creates two sperm and a pollen tube
Ovules/Seeds
- Immature seeds are ovules
- Protective layer of cells: integument
- May be multiple
- Diploid sporophyte tissue
- New structure
- Integument houses megasporangium
- “Nucellus”; fleshier than previous megasporangium
- Nucellus dooes not proliferate; most space after fertilization is the megagametophyte
- One megaspore develops; four developed, but three break down
- Hole in integument: micropyle
- Fertilization/pollen tubes
- After fertilization, it is called a seed
- Zygote develops into embryo; mitosis
- Has a food supply
- Food supply is the megagametophyte
- Integument becomes seed coat
- Micropyle seals off
- Seeds are dormant until good for growth
Gymnnosperms
- Naked seed plants
- Seeds are exposed to air; no fruiting
- Multiple phyla
- Coniferophyta
- Conifers
- Cone-bearing trees
- Representative
Coniferophyta
Reproduction
- Male cones: Strobili
- Relatively small, soft, non-woody
- Only used to make pollen grains
- Leaves (microsporophylls) arrange around a central axis
- Each microsporophyll bears 2 pollen sacs or microsporangia
- Microsporocytes develop via meiosis to make microspores, microspores develop via mitosis to make pollen grains
- Pollen grains are released from cones
- Two prothallial cells: part of the body of the male gametophyte
- Generative cell: Large central cell that develops the two sperm
- Tube cell: develops the pollen tube
- Large air sacs on sides that allow for air dispersal
- Tube cell directs fertilization of pollen tube
- Non-fertilizing sperm breaks down
- Female cones (ovulate cones)
- Central axis is surrounded by cone scales
- Cone scales; bracts
- Bracts are modified leaves
- Cone scales are more than leaves; encompasses the bract and the bud/stem; not a megasporophyll
- Cone scales are under the ovule
- Space between scales allows pollen grain to enter
- Between spore release, scales are shut
- Megasporocyte divides by meiosis to make four megaspores, megaspore divides by mitosis to make a large megagametophyte (thousands of cells)
- Pines have large genomes used in studying
- Winged seeds; wind dispersal
- Also bird or animal dispersal
Diversity
- Pines (Pinus), firs, Douglas-firs, Sequoia, European larch, juniper, cypress, yew
- Mostly isogamous monoecious
- Secondary growth
- Needle-like leaves
- Reduced surface area
- Less vaporization and evaporation
- Pinus bundles
- Tracheids; true vascular tissue
- High importance: Large swaths in northern coniferous forests (Taigas)
- Norway
- Fewer species, but cover large areas of land
Cycadophyta
- Cycads, Sago palms
- Dioecious
- Haustorial pollen tube
- Feeding tube that ruptures open to release sperm in ovule
Ginkgophyta
- Gingko biloba — Maidenhair tree
- Deciduous
- Not evergreen
- Dioecious
- Fleshy, rancid seed coasts with tasty kernals
- Lobed leaves
Gnetophyta
- Welwitschia, Ephedra, Gnetum
- Some have vessels and some lack archegonia
- Not ancestors to angiosperms; convergent evolution
- Dioecious
Angiosperms
- Highly successful
- All have vessels in xylem
- Direct pollination with animals; less waste
- Co-evolution with insect pollinators
- Fruits and their help with seed dispersal
Evolution
- Around for at least 140 million years; Cretaceous
- Aquatic fossils; highly divided leaves and possible floats around leaves
- Earliest likely woody angiosperms
- MRCA between gymnosperms and angiosperms 300mya
- Extinct group from ~225mya: Bennettitales
Basal Lineages
- Amborella
- Woody
- Small reduced flowers
- Water lilies
- Star anise and its relatives
- No vessels
Magnoliids
- Magnolia (M. grandiflora)
Monocots
- All grasses (oats, wheats, corn, barley)
- Palm trees
- Orchids
- Pollen grain has only one opening/aperture for the pollen tube
Eudicots
- Most diverse
- Oak tree flowers (Quercus pyrenica)
- Pollen grain has or was evolved from a grain that has three openings/apertures
Reproductive Cycle
- Stigma holds pollen grain in place
- Pollen grain absorbs water
- Pollen grain tube cell takes signal; forms pollen tube
- Going through transmitting tissue in the style, the placenta, and entering the embryo sac
- Synergids break down and emit attractive chemicals
- Pollen grain enters tube, through synergid, into egg
- One sperm nucleus forms zygote → embryo
- One sperm nucleus fuses with two polar nuclei of central cell → the triploid nucleus of the endosperm
- Ovary ripens into a dry or fleshy fruit
- Dry endosperm or fleshy cotyledons inside the fruit
- Multiple pollen entering during pollination; competitive fertilization
Pollen Grain
- Wall structure has two main parts:
- Exine: Outer structure; highly sculptured and made from sporopollenin
- Intine: Solid continuous layer inner structure; made from cellulose
- Exine is coated with materials from the microsporangium’s tapetum
Outcrossing
- Outcrossing: Breeding between two different organisms
- Vs. Selfing
- Self-Incompatibility methods
- May also increase via
- Differences in timing of maturation (pollen matures faster than stigma, or vice versa)
- Dioecious vs. monoecious development
Self-Incompatibility
- Foolproof Self-Incompatibility
- Rejection of the self’s locus/pollen
- Controlled by alleles at the S locus
- Two forms of self-incompatibility
Gametophytic
- Easier form
- If the pollen grain’s S allele matches either the allele on the stigma or style, a pollen tube will not grow
- Haploid, single allele
- Crop plants: potato/tomato family
- GSI
Sporophytic
- If tapetum materials match the alleles on the stigma or style, the grain will not germinate
- Diploid, two alleles
- Dominant-recessive alterations
- Mustard family
- SSI
Direct Pollination
- Animal pollination
- Pollinators are rewarded by nectar and pollen; pollinator reward
- Nectar: Sugary
- Pollen: Proteinaceous
Pollination Syndromes
- Suites of characteristics evolved to one particular pollinator
- Hummingbirds and columbines with red petals and tubes
- Moths (Darwin)
- Beetles
- Bees and landing platform flowers
- Bats and nocturnal flowers
- Insects and rotting meat flowers
Nectar Guides
- Differences in UV reflection that signal the presence of pollen
Rate of Speciation
- Increased rates of pollination, co-evolution, etc. between pollinators and plants
- Controlling by specific characteristics: bilateral vs radial symmetry
Fruits
- Pericarp: Ovary wall and fruit around the carpal
- Ovary wall develops into the pericarp around the seeds
- Aid in seed dispersal
- Buoyancy in coconut fruits: Water dispersal
- Winged fruits: dandelion seeds and maple fruits
- Wind or animal pollination: Tumbleweed, Puncture vines
- Ant food bodies, feces
- High effort-energy in sexual reproduction
Sexual and Asexual Reproduction
- Vegetative reproduction (Asexual reproduction): Potatoes, strawberries (stolons), Kalanchoe plantlets, aspens
- Sexual reproduction: Farther dispersal
Apomixis
- Reproduction method
- Flowers and fruits used; no fertilization
- One common method:
- Diploid cell and a parent becoming a zygote
- Dandelion
- Non-genetically diverse offspring
Ecology
Global Ecology
Landscape Ecology
- Larger patterns found in ecosystems
- Fire in chaparral and other habitats
- Newer area of ecology; 30-35 years
Ecosystem Ecology
Population Ecology
- Breeding, evolution, intra- and inter-species interactions
- Population characteristics:
- Populations are defined by natural or cultural geographical boundaries
- N refers to population size (in number of individuals)
- Population density
- Dispersion patterns
Organismal Ecology
- Adaptations in various habitats
Areas of Population Ecology
Demography
- Population size and growth; demographics
- Births and immigration vs. deaths and emigration
Life tables
- Age-specific summaries of survival and reproduction
- Cohort grouping: A group of individuals across age classes
- Static grouping: Snapshots of age classes
Survivorship curves
Late Loss
- High risk at old age
- High K selection
- Humans
Early Loss
- High risk at early age
- High r selection
- High birth, seed dispersal
- Oysters
Constant Loss
- No high risk age stage
- Belding’s ground squirrel, hydra
Stair Steps
- Repeat risk stages
- Molting
- Spiders
Population Growth Models
- Omitting immigration and emigration
- Boom and bust growth
- Logistic growth as normal; bust is the byproduct of density-independent factors
- Logistic growth
- Density-dependent factors include territoriality (a subset of limited resources regarding space and/or mates)
Population Cycles
Life History
- All of the traits that affect an organism’s fitness
- How many times an organism reproduces in its lifetime
- Iteroparous: Many iterations
- May take a while to reach reproductive age
- High likelihood of surviving for more than one year
- Oak trees, bumper crops, insects (monarch butterfly)
- Semelparous: Single iteration
- Desert annuals
- SALMON, agave: “big bang reproduction”
- Clutch/Brood Size; number of offspring produced at one time
- Age of sexual maturity/age of first reproduction
- Earlier maturity based on life expectancy (annuals vs. perennials)
- Parental care or offspring quality and size
- Co-adapted traits: A suite of life history traits shared by a population
Weedy or Opportunistic Species
- Advantageous plants that compete for space by growing quickly in open areas
- r-selected species
- Semelparous
- Many young
- Small young and little parental care
- Low survival rates
Equilibrium Species
- k-selected species
- Iteroparous
- Few young
- Larger young and more parental care
- High survival rates
Life-History Continuum
- High fecundity and low survivorship vs. low fecundity and high survivorship
- A population of local populations that are connected by migration
- Local populations may go extinct, but the populations are restored/repopulated via emigration from another population