Lecture #4 – Plant Structure, Growth And Development Image – the Angel Oak 1 Key Concepts: • • • • • • • • What is a kingdom? Why study plants? What makes a plant a plant? The hierarchy of structure – plant cells, tissues and organs Growth Primary growth – elongation Secondary growth – diameter expansion Morphogenesis occurs during growth 2 Carolus Linnaeus (1707-1778) Image – Linnaeus The founder of modern taxonomy defined kingdoms by morphological similarity 3 Linnaeus’ Taxonomic Hierarchy Taxonomic Category Example (taxon) Kingdom Plantae, also Metaphyta = all plants Division (phylum) Magnoliophyta = all angiosperms Class Liliopsida = all monocots Order Asparagales = related families (Orchidaceae, Iridaceae, etc) Family Orchidaceae = related genera (Platanthera, Spiranthes, etc) Genus Platanthera = related species (P. ciliaris, P. integra, etc) Specific name/epithet ciliaris = one species 4 Linnaeus’ Taxonomic Hierarchy Taxonomic Category Example (taxon) Kingdom Plantae, also Metaphyta = all plants Division (phylum) Magnoliophyta = all angiosperms Class Liliopsida = all monocots Order Asparagales = related families (Orchidaceae, Iridaceae, etc) Family Orchidaceae = related genera (Platanthera, Spiranthes, etc) Genus Platanthera = related species (P. ciliaris, P. integra, etc) Specific name/epithet ciliaris = one species 5 Images – the yellow fringed orchid 6 Platanthera ciliaris Linnaeus recognized only 2 kingdoms • If it moved – animal; if it didn’t – plant • Fungi were lumped with plants • The microscopic world was largely unknown Images – the 3 multicellular kingdoms, animals, fungi and plants 7 The 5 kingdom system – developed in the 1960’s and used until recently Diagram – the 5 kingdom system 8 Molecular data supports 3 domain classification scheme Diagram – 3 domain system of classification Kingdoms are defined by monophyletic lineage 9 Classification is Dynamic! Diagram – transition from 5 kingdom to 3 domain system indicating dynamic nature of classification Multicellular eukaryotes remain fairly well defined – the plants, fungi and animals. Classification of single celled organisms is still underway. 10 Current Taxonomic Hierarchy Taxonomic Category Example (taxon) Domain Eukarya = all eukaryotic organisms Kingdom Plantae, also Metaphyta = all plants Division (phylum) Magnoliophyta = all angiosperms Class Liliopsida = all monocots Order Asparagales = related families (Orchidaceae, Iridaceae, etc) Family Orchidaceae = related genera (Platanthera, Spiranthes, etc) Genus Platanthera = related species (P. ciliaris, P. integra, etc) Specific name/epithet ciliaris = one species 11 Why Plants? 12 Why Plants? Image – shooting stars 13 What makes a plant a plant??? 14 Images and diagrams – characteristics that separate plants from other kingdoms 15 What makes a plant a plant??? • Multicellular, eukaryotic organisms with extensive specialization • Almost all are photosynthetic, with chloroplasts (= green) Some obtain additional nutrition through parasitism or carnivory Some are saprophytic, entirely without chlorophyll (absorb dead OM) • Excess carbohydrates stored as starch (coiled, branched polymer of glucose) • Cell walls of cellulose = fibrous (not branched) polysaccharide = accounts for the relative rigidity of the cell wall • Cell division by formation of cell plate • Most extant plant species are terrestrial (many characteristics that are adapted for terrestrial life) • Separated from cyanobacteria by chloroplasts • Separated from green algae by various adaptations to 16 terrestrial life Read this later…. Plants were the first organisms to move onto land • Occurred about 475mya • Very different conditions from former marine habitat • Many new traits emerged in adaptation to life on dry land • Extensive adaptive radiation into many new ecological niches 17 Four major groups of plants have emerged since plants took to land Diagram – phylogeny of land plants; same on next slide 18 We will focus on angiosperms Next semester in 211 you will learn more about the transition from water to land, and the evolution of reproductive strategies in all plants 19 Angiosperms – the flowering plants: 90% of the Earth’s modern flora Images – flowering plants 20 Basic Structure of the Plant Cell – what’s unique??? Diagram – plant cell; same on next slide 21 Basic Structure of the Plant Cell 22 Critical Thinking • Do all plant cells have chloroplasts??? • How can you tell??? 23 Critical Thinking • Do all plant cells have chloroplasts??? • How can you tell??? 24 Critical Thinking • Do all plant cells have chloroplasts??? • How can you tell??? Image – chloroplast free white bracts on white-top sedge 25 More on the cell wall: • All cell walls are produced by the cell membrane, outside • Primary wall is produced first Diagram – primary and secondary cell walls; same on next slide Mostly cellulose • Secondary walls are produced later Lignified, so ??? • Secondary walls are interior to primary walls 26 More on the cell wall: • All cell walls are produced by the cell membrane • Primary wall is produced first Mostly cellulose • Secondary walls are produced later Lignified, so • Secondary walls are interior to primary walls 27 Five Major Plant Cell Types Micrographs – plant cell types • Parenchyma • Collenchyma • Sclerenchyma • Xylem elements • Phloem elements 28 Parenchyma • • • • Thin primary wall No secondary wall Many metabolic and storage functions Bulk of the plant body Micrographs – parenchyma cells 29 Collenchyma • Thick primary wall • No secondary wall Micrograph – collenchyma cells; same on next slide Implications??? • Support growing tissues 30 Collenchyma • Thick primary wall • No secondary wall Implications??? • Support growing tissues 31 Sclerenchyma • Thick secondary wall • Secondary walls are lignified Micrograph – sclerenchma cells; same on next slide Implications??? • Support mature plant parts • Often dead at maturity 32 Sclerenchyma • Thick secondary wall • Secondary walls are lignified Implications??? • Support mature plant parts • Often dead at maturity 33 Collenchyma vs. Sclerenchyma • • • • Both provide structural support Both have thick walls Collenchyma = thick primary wall, no lignin Sclerenchyma = thick secondary wall, lignified Micrographs – collenchyma and sclerenchyma cell comparison 34 Xylem Elements • Lignified secondary walls • Always dead at maturity (open) • Function to transport water and dissolved nutrients, and to support the plant • Tracheids and vessel elements Diagrams and micrograph – tracheids and vessel elements 35 Critical Thinking • Vessel elements and the convergent evolution of rings • What else looks like this???? • What is the function???? Micrograph – rings of lignin in developing vessel element; same on next slide 36 Critical Thinking • Vessel elements and the convergent evolution of rings • What else looks like this???? • What is the function???? 37 Phloem Elements • Sieve tube members + companion cells • STM lack nucleus, ribosomes – their metabolism is controlled by the companion cells Micrograph – phloem elements • Function to transport the products of metabolism • Non-angiosperms have more primitive phloem elements 38 Critical Thinking • What might be the functional advantage of a cell with no nucleus??? Diagram – phloem elements 39 Critical Thinking • What might be the functional advantage of a cell with no nucleus??? 40 Plants are Simple Only Five Major Cell Types Micrographs – plant cell types • Parenchyma • Collenchyma • Sclerenchyma • Xylem elements • Phloem elements 41 Tissue Systems • • • • Diagram – plant tissue types Epidermis Vascular Ground Meristem 42 Epidermis Tissue: • Covers the outer surface of all plant parts • Shoot surfaces covered with waxy cuticle Micrograph and diagram – epidermis Helps to protect the plant and prevent desiccation • Usually a single, transparent cell layer • Tight joints; stomata allow for gas exchange 43 Critical Thinking • Do roots have a waxy cuticle??? • Why or why not??? 44 Critical Thinking • Do roots have a waxy cuticle??? • Why or why not??? Never forget the importance of natural selection!!!!! 45 Vascular Tissue: • Transports water, solutes, and metabolic products throughout the plant • Confers structural support • Includes xylem elements, phloem elements, parenchyma and sclerenchyma fibers Micrograph – vascular bundle in cross section 46 Critical Thinking • Why does vascular tissue give structural support to a plant??? 47 Critical Thinking • Why does vascular tissue give structural support to a plant??? 48 Ground Tissue: • Bulk of the plant body – pith, cortex and mesophyll • Mostly parenchyma • Most metabolic, structural and storage functions Micrograph and diagram – ground tissues in stems and leaves 49 Critical Thinking • Is this what the inside of a tree looks like??? Micrograph – herbaceous dicot stem 50 Critical Thinking • Is this what the inside of a tree looks like??? Micrograph of herbaceous eudicot stem; image of woody stem; diagram of woody stem tissue organization 51 Meristem Tissue: • How the plant grows • Cells divide constantly during the growing season to make new tissues • More details later Image – new growth at tip of stem 52 Plants are Simple Only Four Major Tissue Types • • • • Diagram – plant tissue systems Epidermis Vascular Ground Meristem 53 Tissues Make Organs: • Roots – anchor the plant, absorb water and nutrients • Stems – support the leaves • Leaves – main site of photosynthesis • Reproductive organs (flowers, cones, etc – more later) All organs have additional functions – hormone synthesis, transport, etc… 54 Plant Organ Systems Diagram – root and shoot systems 55 Modern molecular evidence indicates four classes of angiosperms paleoherbs magnoliids eudicots monocots ancestral 56 Paleoherbs and Magnoliids comprise about 3% of angiosperms Paleoherbs • Aristolochiaceae, Nymphaeaceae, etc Magnoliids • Magnoliaceae, Lauraceae, nutmeg, black pepper, etc Images – water lily and magnolia 57 Modern evidence indicates 4 classes of angiosperms paleoherbs magnoliids eudicots monocots ~ 97% of angiosperms ancestral 58 Monocots include grasses, sedges, iris, orchids, lilies, palms, etc….. Images – monocots 59 Eudicots include 70+% of all angiosperms: • Most broadleaf trees and shrubs • Most fruit and vegetable crops • Most herbaceous flowering plants Images – eudicots 60 Monocots vs. Eudicots Monocots • Flower parts in multiples of 3 • Parallel leaf venation • Single cotyledon • Vascular bundles in complex arrangement • ~90,000 species Eudicots • Flower parts in multiples of 4 or 5 • Netted leaf venation • Two cotyledons • Vascular bundles in a ring around the stem • Modern classification indicates 2 small primitive groups + eudicots • 200,000+ species 61 Root System Tissue Organization Eudicots Monocots Micrographs – cross sections of eudicot and moncot roots; same on next 3 slides Epidermis, ground, endodermis, pericycle, vascular tissues 62 Eudicot root – closeup Epidermis Cortex Endodermis Pericycle Vascular tissues – in solid core 63 Monocot root – closeup Epidermis Cortex Endodermis Pericycle Vascular tissues – in ring Pith in the very center 64 Critical Thinking • Where do branch roots form??? 65 Critical Thinking • Where do branch roots form??? Micrograph – root emerging from pericycle 66 Stem System Tissue Organization Eudicots Monocots Micrograph – eudicot and monocot stem tissue organization; same on next 4 slides Epidermis, ground, vascular tissues 67 Eudicot stem – closeup Epidermis Cortex Vascular tissues – bundles in a ring Pith 68 Monocot stem – closeup Epidermis Cortex Vascular tissues – bundles are scattered 69 Wood forms from a meristem that links the vascular bundles: 70 Stem System Tissue Organization Eudicots Monocots Monocots cannot make wood More on wood formation later 71 Leaf Tissue Arrangement Micrograph – cross-section of leaf tissue arrangement Epidermis, ground, vascular tissues 72 Leaf closeup Epidermis Diagram – leaf tissue arrangement Cortex – palisade mesophyll Cortex – spongy mesophyll Vascular tissues 73 Stomata – pores to allow for gas exchange and transpiration Micrograph – epidermis tissue showing stomata 74 See, plants really are simple • 5 cell types • 4 tissue types • 4 organ types Diagram – shoot and root systems 75 Plant Growth • Remember, most plants are anchored by roots • They can’t move to escape or take advantage of changes in their environment • Plants adjust to their environment • Simple structure + lots of developmental flexibility allow plants to alter when and how they grow Developmental flexibility comes from meristems 76 Meristem Tissues • Actively dividing cells that generate all other cells in the plant body • Cause indeterminate growth Stems and roots elongate throughout the plant’s life (indeterminate primary growth) Trees continually expand in diameter (indeterminate secondary growth) Branches form in roots and stems 77 Not all plant parts have indeterminate growth patterns Indeterminate: Roots and Stems Determinate: Leaves Flowers Fruits These parts grow throughout the life of the plant, exploring new environments or responding to damage These parts grow to a genetically +/predetermined size and shape and then stop – cannot repair 78 damage Some mature cells can de-differentiate to become meristematic once more!!! • Primarily occurs in the indeterminate parts Stems and roots • A process that very seldom occurs in other kingdoms • Allows stems and roots to repair damage and form branches and sprouts 79 Critical Thinking • Can all plant cells de-differentiate??? • What would control this??? 80 Critical Thinking • Can all plant cells de-differentiate??? • What would control this??? 81 Critical Thinking • Can all plant cells de-differentiate??? • What would control this??? 82 Growth in Plants: an irreversible increase in size due to metabolic processes (processes that use ATP energy) • Cell division produces new cells = function of meristem • Cell expansion increases the size of the new cells = up to 80% of size increase • Cell differentiation occurs during and after expansion 83 The plane of cell division contributes to morphogenesis Diagram – planes of cell division and the effect on morphogenesis Division in 2 planes forms sheets of cells 84 Critical Thinking • What tissues are files of cells??? • What tissues are sheets of cells??? • What tissues are 3-D bulky??? 85 Critical Thinking • What tissues are files of cells??? • What tissues are sheets of cells??? • What tissues are 3-D bulky??? 86 Growth in Plants: an irreversible increase in size due to metabolic processes (processes that use ATP energy) • Cell division produces new cells = function of meristem • Cell expansion increases the size of the new cells = up to 80% of size increase • Cell differentiation occurs during and after expansion 87 Auxin-mediated cell expansion Diagram – how auxin works to promote cell expansion ATP is used 88 The direction of cell expansion depends on cellulose orientation, and contributes to morphogenesis Diagram – cellulose orientation in primary wall and the effects on morphogenesis 89 Growth in Plants: an irreversible increase in size due to metabolic processes (processes that use ATP energy) • Cell division produces new cells = function of meristem • Cell expansion increases the size of the new cells = up to 80% of size increase • Cell differentiation occurs during and after expansion 90 Expansion and differentiation occur in an overlapping zone in all plant parts Diagram – patterns of growth in roots 91 REVIEW: Growth in Plants: an irreversible increase in size due to metabolic processes (processes that use ATP energy) • Cell division produces new cells = function of meristem • Cell expansion increases the size of the new cells = up to 80% of size increase • Cell differentiation occurs during and after expansion 92 Location of the meristems determines the pattern of plant growth Diagram – location of meristems on the plant body; next slide also Most common meristems: apical, axillary and lateral 93 Apical meristems cause elongation of roots and stems 94 Micrograph – longitudinal section showing distribution of tissues in root 95 Images – root cap and mucigel 96 Root Cap • Protects the meristem • Secretes mucigel Eases movement of roots through soil Secretes chemicals that enhance nutrient uptake • Constantly shedding cells Mechanical abrasion as roots grow through soil • Constantly being replenished by meristem 97 Primary Growth in Roots Diagram – longitudinal section of root showing zones of growth; same on next 2 slides 98 Primary Growth in Roots 99 Primary Growth in Roots 100 Root Hairs • Form as the epidermis fully differentiates • Extensions off epidermal cells NOT files of cells Part of an epidermal cell Micrograph – root hairs extending from epidermis; same on next few slides • Hugely increase the surface area of the epidermis • 10 cubic cm (double handful) of soil might contain 1 m of plant roots Mostly root hairs 101 Critical Thinking • What is the selective advantage of root hairs??? 102 Critical Thinking • What is the selective advantage of root hairs??? 103 Root Hairs • By contrast, 10 cc of soil may contain up to 1000 m of fungal hyphae (1km!) These serve a similar function for the fungus Ramify throughout the substrate for maximum absorption Some fungi form symbiotic associations with plant roots and both organisms benefit from this huge absorptive surface area! More in 211….. 104 Apical meristems cause elongation of roots and stems Diagram – location of apical meristems 105 Apical Meristems in Shoots Micrograph – longitudinal section of stem showing apical and axillary meristems 106 Critical Thinking • There is no “shoot cap” – why not??? 107 Critical Thinking • There is no “shoot cap” – why not??? 108 Axillary meristems allow for branching – similar in structure and function to apical meristems Diagram – meristem locations Remember, pericycle in roots has same function 109 Axillary Meristems in Shoots Micrograph – longitudinal section of stem showing apical and axillary meristems; same on next two slides 110 Primary Growth in Shoots • Apical meristem • Leaf primordia • Axillary buds 111 As with roots – cell division occurs first; zones of expansion and differentiation overlap Axillary buds may activate to make branches, or may remain dormant 112 Primary growth of a shoot – elongation from the tip Diagram – how stems elongate during primary growth 113 Lateral meristems cause diameter expansion Diagram – meristem locations Roots also expand in diameter, but it’s more complicated – we’ll save that for BIOL 300 114 Lateral Meristems = Cambiums Diagram – lateral meristems 115 Remember: Diagram – primary vs. secondary growth Elongation is primary growth Diameter expansion is secondary growth 116 Secondary growth – diameter expansion Images – cross section of wood and whole tree 117 Eudicot Stem – recall the arrangement of vascular bundles Micrograph – cross section of a eudicot stem; same on next 2 slides 118 Eudicot Stem – recall the arrangement of vascular bundles Vascular cambium forms here: 119 Eudicot Stem – recall the arrangement of vascular bundles Vascular cambium forms here: a cylinder of meristem tissue between the xylem to the interior and the phloem to the exterior 120 Secondary xylem and phloem form through cell division by the vascular cambium Diagram – location of the vascular cambium relative to other tree tissues 121 During primary growth the vascular tissues form in bundles from the apical meristem Diagram – transition from primary growth to secondary growth; same on next slide During secondary growth the vascular tissues form in cylinders from the vascular cambium 2o xylem to the inside 2o phloem to the outside 122 Secondary xylem accumulates 123 Secondary Xylem = Wood! Micrograph – cross section of woody plant showing secondary tissues; same on next slide 124 Annual growth rings are accumulating rings of secondary xylem 125 Critical Thinking • Why do eudicot trees taper??? Diagram – pattern of accumulation of secondary xylem as a tree grows; same on next slide 126 Critical Thinking • Why do eudicot trees taper??? 127 Bark • All tissues external to the vascular cambium • Diameter expansion splits original epidermis Bark structurally and functionally replaces epidermis • Inner bark Functional secondary phloem • Outer bark Composition varies as tree matures 128 Bark Formation Micrograph – cross section of a tree showing bark formation 129 Cork Cambium • Meristematic tissue • Forms in a cylinder during 2o growth • Divides to produce cork cells Cells filled with waxy, waterproof suberin • Eventually cork cambium becomes cork itself 130 More on cork cambium • First layer develops from cortex De-differentiation!!! • Second layer forms from cortex – same process • Third layer forms from cortex….. • Cortex eventually runs out • Then what??? 131 More on cork cambium • First layer develops from cortex De-differentiation!!! • Second layer forms from cortex – same process • Third layer forms from cortex….. • Cortex eventually runs out • Then what??? 132 More on cork cambium • First layer develops from cortex De-differentiation!!! • Second layer forms from cortex – same process • Third layer forms from cortex….. • Cortex eventually runs out • Then what??? 133 More on cork cambium • First layer develops from cortex De-differentiation!!! • Second layer forms from cortex – same process • Third layer forms from cortex….. • Cortex eventually runs out • Then what??? 134 Critical Thinking • What is the next available layer of tissue??? Diagram – lateral meristems and the secondary tissues in a tree; same on next slide 135 Critical Thinking • What is the next available layer of tissue??? 136 More on cork cambium • First layer develops from cortex De-differentiation!!! • Second layer forms from cortex – same process • Third layer forms from cortex….. • Cortex eventually runs out • Then what??? 137 More on cork cambium • First layer develops from cortex De-differentiation!!! • Second layer forms from cortex – same process • Third layer forms from cortex….. • Cortex eventually runs out • Then what??? 138 Stem Tissue Derivations and Fates: Diagram – how undifferentiated cells develop into the tissues of the plant body Cells divide, expand and differentiate 139 Review: Key Concepts: • • • • • • • • What is a kingdom? Why study plants? What makes a plant a plant? The hierarchy of structure – plant cells, tissues and organs Growth Primary growth – elongation Secondary growth – diameter expansion Morphogenesis occurs during growth 140 Monocots, Palmetto Trees, Ft. Moultrie and the SC State Flag Various images and a micrograph of a monocot stem – an example of one influence of plants on American history 141