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Switching gears from algae to angiosperms Seagrasses blade holdfast flower leaves stem roots/rhizomes • Less tissue specialization • Happy in salt water • More tissue specialization • Stressed by salt water 1 2 Algal Evolution: Angiosperm Characteristics 3.9 bya = Cyanobacteria appear and introduce photosynthesis 2.5 bya = Eukaryotes appeared (nuclear envelope and ER thought to come 1) Pigments: chl a &chl b cartenoids- B-carotene, violaxanthin from invagination of plasma membrane) 1.6 bya = Multicellular algae -Rhodophyta (Red algae) &Chlorophyta (Green algae) 2) Chloroplast structure: 2membranes stacks of 2-6 3) Storage product: starches:amylose & amlyopectin 900 mya= Dinoflagellates & Invertebrates appear 4) Flagella: None depend of pollinators, wind, birds, bugs, bats 5) Mitosis: double fertilization sperm cell undergoes mitosis 1 sperm fuses with egg to form zygote 1 sperm fuses with polar nuclei to form endosperm - nutritional source for growing zygote 490 mya = Phaeophyceae (Brown algae) & land plants & coralline algae & crustaceans & mulluscs 408mya= Insects & Fish 362 mya = Coccolithophores & Amphibians & Reptiles 290mya- Gymnosperms 3 145 mya = Diatoms & Angiosperms 1 Groups (Kingdom) DOMAIN 1.Bacteria- cyanobacteria (blue green algae) 2.Archae 3.Eukaryotes 1. Alveolates- dinoflagellates Unicellular, freshwater Chloroplast peptidoglycan 2. Stramenopiles- diatoms, heterokonyophyta Plantae 3. Rhizaria- unicellular amoeboids Rhodophyta Chlorophytes phycoerythrin 4. Excavates- unicellular flagellates 5. Plantae- rhodophyta, chlorophyta, seagrasses 6. Amoebozoans- slimemolds 7. Fungi- heterotrophs with extracellular digestion Chl b, Starchamylose & amlyopectin Glaucophytes Charophytes Embryo, cuticle Land Plants 8. Choanoflagellates- unicellular 9. Animals- multicellular heterotrophs 5 6 Adapted from Sadava 2014 Types of flowering plants 1. Seagrass taxonomy Mesophytes/ Glycophytes- grow where freshwater is available & lack specialized adaptations that prevent water loss 2. Hydrophytes- live in water, partially or fully submerged (seagrass) 3. Xerophytes- have, morphological, anatomical, & reproductive adaptations to aid in the retention of water ( mangroves & salt marsh plants) 1. Halophytes- adaptations to prevent water loss & can grow in saline habitats 1. Facultative- do not require saline conditions 2. Obligate- specific requirement for sodium to complete their life cycle 7 Domain Eukaryote Kingdom/Clade Plantae Phylum/Division Magnoliophyta - angiosperms Class Liliopsida - monocots Order Najadales y Zosteracea- eel g grass Family Genus Phyllospadix- surf grass species scouleri 8 2 Evolution of Seagrasses Seagrasses are Monocots Monocots - one cotelydon (embryotic leaf) - flowers petals either 3, 6, 9 - leaves with parallel blades more closely related to lilies than grasses - ~60 spp; present on all continents except Antarctica. - Tropical and temperate spp; shallow, soft sediment nearshore environments - Initially, debate on whether seagrasses evolved from coastal plants (salt marsh or mangroves) or freshwater plants 9 Seagrass Distribution - Recent molecular work suggests both are correct: polyphyletic origins of seagrasses ( at least three evolutions of salt water habitat) 10 SEAGRASS 1. Stabilize sediments, prevent erosion - physically & chemically change the sediment through release of O2, decomposition of subterranean parts, bioturbidation through growth of roots & rhizomes 2. Improve water clarity by trapping particulates 3. Seagrass meadows = incredibly productive environments - High Ps rate; 2000-4000 g C/m2/yr; on same order as sugar cane -Importance of detrital food web; epiphyte productivity; use of stable isotope analysis to identify both - 2ndary productivity- biomass of heterotrophic organisms in a system 11 12 3 4. Nursery role for many commercially important fish and inverts Adaptations to marine life Hydrophytes-Ability to live completely submerged in salt water - No stomates (osmoregulation and CO2 uptake through epidermal cells) - Chloroplasts only in thin layer of epidermal cells 5. Important trophic linkages to other ecosystems - Hydrophilic pollen - Bladelike morphology with sheath high energy environments - Lots of things eat in one place, poop/die in another - Lacunae & aerenchyma = air spaces e.g., grunts in the Caribbean; hide out during the day in coral crevices, forage at night in seagrass meadows - Seagrass leaves 13 14 Biomass & Productivity Seagrass morphology Below-ground Biomass- 50 -90% of standing stock Upper Leaf Blades- photosynthesis Basal Leaf Sheaths- colorless, covers short shoot and blades, protects the apical meristem Short Shoots- erect stems that produce flowers and leaves, not involved in vegatative reproduction Rhizome = underground primary stem, storage of starch; transport of photosynthate, nutrients, both horizontal and vertical – physiological integration of beds. Leaf Area Indices- amount of leaf available for photosynthesis expressed relative to ground cover broad leaf trees- 4-6 grasses- 9-10 rainforest leaves- 20 seagrasses- 4.9-21 4 9 21 LAI of seagrass reflect the high density of blade cover in relation to ground cover, adaptation to submarine irradiance, need for water movement to enhance nutrient and gas diffusion Roots = adventitious, thick to fibrous, have root cap & root hairs 15 16 4 Seagrass morphology Seagrass Leaf Diagram Bundle sheath- containing phloem & xylem lacunae aerenchyma Leaves - flat and flexible -strong due to internal fiber bundle -reduced vascular bundle with xylem and phloem - epidermal cells lack stomata & guard cells - covered by a thin porous cuticle Fb b Fiber bundles dl Wheat Bundle sheath- containing phloem & xylem stomata lacunae aerenchyma 18 Fiber bundles 17 Seagrass morphology Seagrass morphology Lacunae- gas exchange - air spaces formed by the breakdown of cells - continuous through the blades, short shoots, rhizomes, & roots - provide gas exchange through plant - storage of CO2 - diffusion of O2 into the rhizome & roots - have septae (diaphrams) with pores allow air exchange but block water to prevent flooding Aerenchyma tissue- gas exchange - Formed by cell separation - Specialized parenchyma tissue with air spaces - Mechanism M h ism for f roott aeration ti iin llow oxygen concentrations 19 20 5 Seagrass Habitat Requirements Seagrass Reproduction - Shallow water (above 20m) - 2 kinds of reproduction: sexual and clonal - Genets – (genetic individual) consists of a series of ramets (physical individual)- short shoots - majority dioecious whereas only 5% of all angiosperms dioecious - Very susceptible to turbidity require clear water - Higher Ec (photosynthesis = respiration) than algae due to below ground biomass - Lower photosynthetic capacity than terrestrial p plants - Less carbon available for Ps in water than in air - Macroalgae are better at using HC03- than seagrasses. - When seagrasses evolved higher atmospheric C02, more inorganic C02 available in sea water. - Seagrasses have lower nutrient requirements than algae 21 Seagrass Reproduction 22 Seagrass Reproduction Clonal- ability to carry out vegetative expansion via a rhizome that produces short shoots or ramets - sharing of resources btw ramets in areas of stress or low nutrients • Zig-zag pattern of flowers in the spadix (spadices) • Male flowers release long, stringy pollen (~2 mm long!) in clumps, typically on an incoming i i tide, dispersed by water motion Hydrophily = water-mediated pollination Posidona meadows in Mediterranean and Australia - one individual may cover 10 m2! - Genetic work to ID individual genets 23 Anthers and pollen 24 6 Seagrass Reproduction Seeds developing in spadix Vivipary- bearing live young - germination occurs while seed is still attached to flower - mangroves also do this 25 Seagrass Reproduction • Fertilization occurs between pollen and stigma of female flowers, both are coated with a strongly adhesive substance 26 Seagrass Reproduction • Sexual reproduction not nearly as common as clonal • Often high rates of seed predation • Positive density-dependence (facilitation by adult surfgrass) and facilitation by coralline alga for recruitment mm • Ripe fruits are released from the spadix; pericarp erodes away, leaves two bristly arms 27 28 7 Shelton Ecology 200830 29 Shelton Ecology 2008 Surfgrasses Zostera marina eel grass • Monoecious • Mediterranean & Temperate • Common locally in bays and estuaries • Wide blades (3-10mm) • Protogynous (female first, then male) Thalassia testudinum Phyllospadix scouleri turtle grass • Dioecious • Tropical – common in Caribbean • Food for sea turtles, manatees, dugongs • roots for nutrient uptake 31 Flatter, wider blades (2-4 mm) • Higher, lower wave action Phyllospadix torreyi Rounder, thinner blades (1-3 mm) • Lower, higher wave action • Dioecious, long-lived perennial • Only genus to occur on rocky substrate •roots for attachment only 32 8 Surfgrass Removal Experiments Removal pools 10-150 C higher Temperature difference are biologically significant 33 34Shelton 2009 Seagrass Community Ecology Seagrass Community Ecology Lots of grazers, especially in the tropics epiphytes Urchins, manatees, dugongs, green turtles Smithora Water fowl, fish 35 36 9 Seagrass Community Ecology Major (ongoing) loss of seagrass habitat worldwide associated mesograzers- feed on associated epiphytic algae declines reported since 1970 from: Heminga and Duarta, 2000 anthropogenic natural 37 38 Seagrasses Habitat Restoration Major (ongoing) loss of seagrass habitat worldwide Mitigation for seagrass loss: - Protected under Clean Water Act & Magneson-Stevens Act (fisheries focus, “essential fish habitat”) 1) Coastal development: dredging and filling 2) Eutrophication 3)) Oilspills p 4) Thermal pollution from power plants 5) Invasive species – Caulerpa taxifolia 6) Anchors & prop scars - Test of ecological understanding: SEAGRASS RESTORATION 39 40 10 Caveats to doom and gloom scenario? Things we’ve learned so far: 1. Building a seagrass meadow is really hard Historical losses: “wasting disease” on East Coast, 1931-1950s 2. We don’t even really have a good metric for success Blackish-brown spots, loss of leaves. ~90% SG lost on Atlantic coast, within a year or so 3. Subtle things may make a big difference in “quality” of a seagrass bed e.g. Genetic G ti diversity di it of f ttransplanted l t d vs. natural t l seagrass meadows d Unclear what caused death or recovery (Labyrinthula slime mold? Bacteria? Ascomycete fungus? Temperature spike?) Did fisheries collapse? Not really. 41 42 11