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Lecture #7 Angiosperms: Form & Function Plant cells • contain all the usual eukaryotic “suspects” – mitochondria, Golgi, ER, vacuoles etc…. – double phospholipid plasma membrane with embedded proteins and carbohydrates – BUT they also possess a cell wall – cytoplasm and the organelles are sometimes referred to as the protoplasm – nucleus is similar to the animal cell nucleus Plant cells • unlike animal cells – plant cells are capable of storing large quantities of many substances • some substances are stored within the cytoplasm – other materials are stored within vacuoles • primary central vacuole – water and salts and wastes • spherosomes or lipid bodies - lipid • plant cells also possess specialized structures Plastids – plastids – group of dynamic organelles that are able to perform many functions • made of an inner membrane and an outer membrane with a stroma in between • several types within a plant cell • 1. chloroplasts – forms from immature pro-plastids once the cell is exposed to light – contain the photosynthetic pigment chlorophyll and all the accessory proteins and complexes required for photosynthesis – inner membrane is elaborately folded into membrane sheets called thylakoid membranes (large surface area for photosynthetic machinery) – in certain areas the thylakoid membranes is folded into vesicle-like bags stacked together – grana (transport of H+ ions required in photosynthesis) chloroplast Plastids • 2. amyloplasts – in plants tissues that can’t photosynthesize – roots, bark and wood cells – accumulate sugar and store it as starch • 3. chromoplasts – e.g. in tomatoes and yellow squash – bright red, yellow and orange lipids accumulated here • 4. leucoplasts – large and unpigmented plastids – no chlorophyll or lipid pigments – involved in the synthesis of fats and phospholipids potato amyloplasts tomato chromoplasts The Cell Wall • all cells of the plant have cell walls (except the sperm) • it is an active, dynamic organelle with many metabolic functions • contains large amounts of cellulose – in the form of cellulose microfibrils – made by rosette protein complexes • all cells have a thin primary cell wall – made of cellulose – also contains pectin – complex polysaccharide that allows for plant growth – pectin also forms the middle lamina found between two plant cells middle lamina – “cement”-like layer of pectin found between two plant cells The Cell Wall • in cells that require strength – there is a thicker secondary wall that is synthesized from the protoplasm – laid down in between the primary wall and the plasma membrane – the secondary wall is almost always impregnated with lignin – lignin is strong and resists fungal and bacterial attacks both the primary and secondary walls are permanent and are not degraded or depolymerized • plant cells are in communication with one another – despite the presence of the cell wall • connections between cells = plasmodesma (plural = plamsodesmata) – the plasma membrane of one cell passes through the plasmodesma and is continuous with the PM of the adjacent plant cell – often plant cells have clusters of the PDs – called pit fields • there exists a diffusional space from one cell wall to another = apoplast • the transport of materials from one cell to another through this cell wall space = apoplastic • the transport of materials from the inside of one cell through the plasmodesma into the neighbouring cell = symplastic • the presence of plasmodesmata means that the individuality of the plant cell is eliminated • the plant becomes one interconnected mass of cells and protoplasm = symplast Plant Cells • three types of plant cells: • classified based on the nature of their cell walls • 1. parenchyma – only have thin primary walls – undifferentiated cells – differentiation leads to the other two cell types • 2. collenchyma – primary cell walls thin in some areas, thick in others • 3. sclerenchyma – primary and secondary walls containing lignin 1. Parenchyma • only possess thin primary cell walls • these cells comprise parenchyma tissue – ground tissue that fills the gaps in between other tissue types • most common type of cell and tissue within the plant • relatively undifferentiated • metabolically active • most usually remain alive once they mature • capable of dividing even after they mature = known as meristematic 1. Parenchyma • some parenchyma cell subtypes are specialized for specific tasks – a. chlorenchyma cells – parenchymal cells containing chloroplasts – b. glandular cells – secrete – c. transfer cells – mediate short distance transport of material • parenchyma tissues can be classified according to their function palisade – spongy parenchyma – round loosely packed parenchyma cells in the leaf – surround air spaces – palisade parenchyma – columnar shaped parenchyma cells found beneath the epidermis of the leaf spongy 2. Collenchyma collenchyma – thin primary cell wall in some areas • in other areas the cell wall thickens – most often the corners of the cell • cells exhibit plasticity – the ability to become deformed by pressure or tension and to retain this new shape once this force is removed • collenchyma tissue - usually only produced in elongating shoot tips – give the tips strength as it elongates but it can be stretched – found just underneath the epidermis parenchyma sclerenchyma 3. Sclerenchyma – has both primary and secondary cell walls • these cell walls are lignified • have the property of elasticitythe ability to become deformed by pressure or tension and to return to normal shape once this force is removed • most sclerenchyma cells die once they mature • because they only need to provide strength to the plant • some can remain alive and metabolically active 3. Sclerenchyma • two types of sclerenchyma cells – conducting and mechanical – mechanical – comprised of long fibers and short sclereids with thick secondary walls – conducting – makes up vascular tissues • sclerenchyma tissue: develops mainly in organs that have stopped growing and have achieved their final shape • the absorption of water by collenchyma and parenchyma tissues can cause swelling – sclerenchyma tissues prevent the cell from expanding stone cells in “pit” of a pear Plant Tissues • found throughout the plant – i.e. in roots, stems and leaves – so each forms a tissue system that is continuous through all parts of the plant • 1. dermal • 2. vascular • 3. ground • 1. dermal tissue - plant’s outer protective covering – forms the first line of defense – usually a single tissue – epidermis – epidermis = single layer of parenchyma cells – cells are tightly joined together Plant Tissues epidermis epidermis – epidermis • forms specialized structures in roots, stems and leaves – e.g. roots - root hairs – e.g. stems – trichomes – hair like outgrowths from the stem epidermis that make it difficult for insects to land on Plant Tissues • in leaves and most stems – epidermis is covered with a waxy cuticle to prevent water loss & protect against pathogen damage (fungi & bacteria) – cuticle is made of a fatty protein called cutin – in very dry environments – there may be an additional layer of wax • in woody plants – the periderm replaces the epidermis in older regions of the stem and roots Trichromes – epidermis: • outer walls contact the environment – regulate the exchange of materials • BUT the presence of cutin can inhibit the entry of CO2 needed for photosynthesis • epidermis contains pairs of cells (guard cells) that surround a hole in the epidermis (stomatal pore) • guard cells + stomatal pore = stoma (plural = stomata) • guard cells control the size of the pore – control CO2 entry – absorb water and swell – increased turgid pressure curves the cell and opens the pore 2. Vascular tissues • plant vascular system is NOT a circulatory system • vascular tissue: xylem & phloem – xylem: for the conduction of water & minerals • conducting cells: tracheids & vessel elements • water and minerals enter the xylem in the roots and are conducted upward to the leaves and stems – phloem: for the conduction of sugars • conducting cells: sieve cells and sieve tube members • phloem picks up sugar from where it is abundant in the plant and transports it to where it is needed PARENCHYMA CELLS Xylem WATER-CONDUCTING CELLS OF THE XYLEM – two types of conducting cells: tracheids and vessel elements Vessel Tracheids 100 µm • either type of cell can be called a “tracheary element” – both are types of conducting sclerenchyma cells – tracheids and vessel elements develop first as immature parenchyma cells with thin primary walls – cell elongates and deposits and secondary wall – cell then dies and the protoplasm degenerates – leaving a hollow dead cell comprised of two cell walls – now a type of sclerenchyma cell Pits Tracheids and vessels (colorized SEM) Vessel element Vessel elements with perforated end walls pit-pair aligned perforations Tracheids • xylem: – tracheids: long cells with tapered ends • tracheids obtain water from those under them and pass the water on to those above them – connection is known as a pit-pair • pit-pair has a pit membrane – provides some friction to water flow • xylem: – vessel elements: shorter and wider cells with flat ends • posses a large hole at either end of the cell = perforation • the vessel elements align their perforations to form a vessel • very little friction to water movement (vs. tracheid) • while the vessel elements are short in length – the vessels they create can run from root tip to shoot tip vessel element vessel element tracheid hardwoods of angiosperms are mainly vessel elements softwoods of gymnosperms are mostly tracheids PARENCHYMA CELLS • xylem: – many tracheids or vessel elements can be identified by their secondary wall – secondary wall is organized as rings = annular thickenings • the majority of the primary wall is uncovered - for water entry and exit • large surface area for water uptake and movement – the strongest type of xylem cell is called a pitted tracheary element • the majority of the primary wall is covered with secondary wall except for small regions called circular bordered pits • water uptake and movement through these is slow aligned perforations tracheids vessel elements Phloem – two cell types: sieve cells and sieve tube members – either are referred to as sieve elements – are parenchyma cells with only a primary cell wall – as sieve elements develop – the plasmodesmata enlarge and become sieve pores • sieve areas – clusters of sieve pores • sieve plates are two stacked sieve areas SUGAR-CONDUCTING CELLS OF THE PHLOEM Sieve-tube members: longitudinal view (LM) Companion cell Sieve-tube member Plasmodesma Sieve plate Nucleus Cytoplasm Companion cell Sieve-tube members: longitudinal view 30 µm 15 µm Sieve plate with pores (LM) Phloem – sieve cell is long and spindleshaped – sieve tube members are shorter and wider with flat ends – these cells must remain alive for function – yet their nuclei degenerate – sieve tube members are associated with companion cells • still retain a nucleus and provide the sieve cells with the necessary nuclear control over their metabolism – sieve cells are associated with albuminous cells Vascular Bundles • xylem and phloem occur together in roots and stems as vascular bundles • bundles are located just interior to the cortex • the arrangement of the vascular bundles helps define a monocot stem from a eudicot stem • also arranged differently according to whether they are in a root or a stem – e.g. in angiosperms there is a solid central vascular cylinder called a stele 3. Ground Tissue • tissues that are neither dermal or vascular – includes cells specialized for storage, photosynthesis, and support – made up of: • 1. parenchyma – made up of parenchyma cells – many parenchyma cells contain chloroplasts • 2. sclerenchyma – made up of sclerenchyma cells – provides support to the plant • 3. collenchyma- made up of collenchyma cells – found in the cortex and pith of stem, the cortex of the root, the mesophyll of leaves and the endosperm of seeds 3 Plant organs • plants respond to changes in their environment by altering their growth • plants have organs each comprised of specific tissues – organ = made up of multiple tissues • 3 organs – form a root system and a shoot system – 1. stems – for transport & support of leaves – 2. leaves – for photosynthesis – 3. roots – for absorption Reproductive shoot (flower) Terminal bud Node Internode Terminal bud Vegetable shoot Leaf Shoot system Blade Petiole Axillary bud Stem Taproot Lateral roots Root system Stems • 1. organ that raises or separates leaves, exposing them to sunlight • 2. also raises reproductive structures – facilitating the dispersal of pollen and fruit • stem – is the main axis • shoot – stem plus any leaves, flowers or buds off of the stem • stems or shoots consist of: Terminal bud Bud scale Axillary buds This year’s growth (one year old) Leaf scar Node Stem Internode One-year-old s shoot formed from axillary bud near shoot apex – a. an alternating system of nodes – where leaves are attached – b. internodes – between the nodes Last year’s growth (two years old) Leaf scar Scars left by termina bud scales of previou winters Growth of twoLeaf scar years ago (three years old) Stems • the arrangement of leaves on the stem = phyllotaxy – so that two leaves don’t lie over each other and shade one another – 1. opposite - two leaves on opposite sides of the stem at the node – 2. whorled – three or more leaves per node – 3. alternate – leaves alternate up the stem; one leaf per node – 4. spiral – the nodes themselves “spiral” up the plant; the leaves can be opposite, alternate or whorled at the nodes Evolutionary Adaptations of Stems • Rhizomes – a horizontal shoot that grows just below the surface – can give rise to vertical shoots • Bulbs – are vertical underground shoots – consist mainly of enlarged, fleshy leaves that store food • Stolons – horizontal shoots that grow along the surface – often called “runners” – asexual reproduction – plantlets can form along the runner where it encounters a suitable habitat • Tubers – enlarged ends of rhizomes or stolons – specialized for storing food – “eyes” – are clusters of axillary buds that mark the nodes rhizome • the angle formed by a stem and leaf – called an axil – the location of the axillary bud – structure that can form a lateral shoot (e.g. branch stem or a leaf petiole) or a flower – vegetative bud if it forms a shoot – floral bud if it forms a flower – can be covered with thick modified leaves = bud scales – new growth from these buds results in “scars” Apical bud Axillary buds Stems • most stem growth is concentrated near the stem tip – consists of an apical or terminal bud – if the apical bud is near an axillary bud can prevent the growth of a new shoot – apical dominance = inhibition of axillary bud growth by the nearby apical bud – removal of the apical bud can stimulate growth of axillary buds and new shoots • axillary and apical buds contain an undifferentiated tissue called meristem Stems • stems grow longer by creating new cells at their tips • growth is at regions known as shoot apical meristems (SAMs) • growth is via mitosis • as the small daughter cells grow to the size of the parent – they push the meristem upward – lower and older cells mature and become part of the growing stem • region below the apical meristem = subapical meristem – site of differentiation • apical meristem is flanked by small, developing leaf primordia which protect the AM Stem growth: Apical meristems the apical meristem and leaf primordia = bud apical meristem developing vascular tissue subapical meristem developing leaf primordia axillary bud Stem growth: Primary and Secondary growth • primary growth in the AM leads to the formation of the subapical meristem • Subapical Meristem is composed of three types of subapical cells – 1. protoderm – gives rise to the epidermis – 2. provascular tissue – gives rise to primary xylem and primary phloem – 3. ground meristem – gives rise to pith and cortex • primary growth is followed by secondary growth – continued differentiation – 1. epidermis becomes the cork cambium (in some plants) – 2. provascular tissue gives rise to the vascular cambium - becomes the secondary xylem and phloem (woody tissues) – 3. pith becomes the interfasicular cambium and the cortex forms the cork cambium (which forms cork) Stem growth: Primary and Secondary growth • subapical meristem – region where division and differentiation takes place – some subapical meristem cells stop dividing and form the first tracheids and vessel elements protoxylem or primary xylem (first xylem) • the remaining meristem cells keep dividing but eventually stop and differentiate into larger tracheids and elements – called metaxylem Stem growth: Subapical meristem T metaxylem V protoxylem Stem growth: Subapical meristem – on the outer edge of the developing vascular bundle the exterior cells are called protophloem or primary phloem • those cells closest to the metaxylem form metaphloem cells • metaphloem is bigger than primary phloem and lasts longer – known often as just phloem • metaphloem cells are much smaller than metaxylem T metaxylem V protoxylem Stems: Internal organization • epidermis – layer of parenchyma cells covered with a cuticle • covered with a waxy cuticle to prevent water loss & protect against pathogen damage (fungi & bacteria) • cortex – interior to the epidermis – simple and homogenous in most stems – composed of photosynthetic parenchyma and sometimes collenchyma Stems: Internal organization • vascular bundles – xylem and phloem – unique arrangement depending on whether the plant is a eudicot or a monocot • pith – most interior portion of the stem – region of parenchyma – similar to the parenchyma of the cortex Stems: Internal organization • vascular bundles – each bundle has xylem and phloem strands running parallel to each other – vascular bundles first contain both primary xylem and primary phloem • then develop metaxylem and metaphloem – monocots – distributed as a complex network throughout the inner part of the stem • frequently described as “scattered” in arrangement – eudicots and gynmnosperms – vascular bundles are arranged in the periphery • surrounding an inner ground tissue of parenchyma called the pith Phloem Xylem Sclerenchyma (fiber cells) Ground tissue Ground tissue connecting pith to cortex Pith Epidermis Key Cortex Epidermis Vascular bundles Dermal Ground 1 mm A eudicot (sunflower) stem. Vascular bundles form a ring. Ground tissue toward the inside is called pith, and ground tissue toward the outside is called cortex. (LM of transverse section) Vascular Vascular bundles 1 mm A monocot (maize) stem. Vascular bundles are scattered throughout the ground tissue. In such an arrangement, ground tissue is not partitioned into pith and cortex. (LM of transverse section) Medullary Rays • vascular bundles of monocots – between the bundles is parenchyma – frequently described as “scattered” in arrangement – more complex than a random arrangement • vascular bundles of eudicots and gymnosperms – vascular bundles are arranged in the periphery surrounding an inner tissue of parenchyma called the pith (ground tissue) Dicot Monocot Stem Vascular Bundles • cap of sclerenchyma on top of primary phloem and metaphloem – called phloem in the figure • below this is a region of large tracheids & vessel elements = metaxylem – called xylem in the figure • smaller tracheids and vessel elements below this is primary xylem • most interior layer is another layer of sclerenchyma Eudicot Stem Vascular Bundles • cap of sclerenchyma • region of phloem (mostly metaphloem) • below this is a region of fascicular/vascular cambium – play a role in secondary growth of a dicot stem – produces secondary xylem and phloem in the “woody” stem – no VC is mature monocot stems (no secondary growth) Eudicot Stem Vascular Bundles • below this is a region of primary xylem & metaxylem • below these bundles is the parenchyma of the pith • parenchyma between vascular bundles are medullary rays Leaves • the main photosynthetic organ • consist of: – 1. a flattened blade – 2. stalk called the petiole - joins the leaf to the stem at the node • contains vascular tissue in the form of veins – monocots – parallel veins – eudicots – branched network Blade Vein Petiole Axillary bud Leaves • blade morphology: – simple – single undivided blade – compound – blade consists of multiple leaflets – double compound – each leaflet divides into smaller leaflets Simple leaf Petiole Axillary bud Leaflet Compound leaf • like compound leaves – this morphology may resist tearing by strong winds Petiole Axillary bud Doubly compound leaf Leaflet Petiole Axillary bud Evolutionary Adaptations of Leaves Tendrils. • tendrils –modified leaves or lateral branches capable of wrapping around small objects – e.g. pea plants, ivy • spines – non-photosynthetic – e.g. cacti – photosynthesis carried out by the stem • needles – capable of photosynthesis Spines. – usually seen in gymnosperms • storage leaves – adapted to storing water – e.g. succulents • reproductive – leaves that produce adventitious plantlets – e.g. succulents Storage leaves. • bracts – often mistaken for petals; modified leaves that surround a group of flowers – e.g. pointsettia Bracts. Leaf Tissues -leaves are comprised of three tissue types 1. epidermis 2. mesophyll 3. vascular tissue - veins Key Guard cells to labels Dermal Ground Vascular Cuticle Sclerenchyma fibers Stomatal pore Epidermal cells 50 µm Surface view of a spiderwort (Tradescantia) leaf (LM) Stoma Upper epidermis Palisade mesophyll Bundlesheath cell Spongy mesophyll Lower epidermis Guard cells Cuticle Xylem Phloem Cutaway drawing of leaf tissues Vein Guard cells Vein Air spaces Guard cells 100 µm Transverse section of a lilac (Syringa) leaf (LM) Leaf Tissues – 1. epidermis: comprised of a cuticle, guard cells, stomata and trichomes • water loss through the epidermis = transpiration • similar to the stem epidermis – large, flat epidermal cells with guard cells and trichomes interspersed • epidermal cells: contain a coating of cutin and wax • guard cells & stomata: upper and lower epidermis have different characteristics – upper surface has fewer stomata if any at all • trichomes – provide shade to the upper surface (deflects sunlight) and slow water loss through the stomata on the lower surface • also make it difficult to eat the leaf by insects Leaf Tissues – 2. mesophyll: ground tissues of the leaf located just below the epidermis • upper surface is a layer of cells called the palisade parenchyma – main photosynthetic tissue of the plant- usually one layer thick – BUT can be several layers thick in regions of intense sunlight • layer below is made of spongy mesophyll – loosely packed parenchyma cells with many intercellular air spaces to permit diffusion of CO2 toward the palisade cells – for photosynthesis vascular bundles • 3. vascular tissues or veins – located between the palisade and mesophyll layers – in dicots – branched venation – large midrib (or midvein) and several smaller lateral veins with narrower branches called minor veins – primary xylem on the upper side and primary phloem on the lower side – vascular tissue of larger veins is surrounded by a bundle sheath (made of bundle sheath cells) collenchyma DICOT LEAF • 3. vascular tissues or veins – monocot – parallel venation • distinct pattern of phloem and xylem • xylem with a few very large vessel elements (3 of them) bundle sheath Roots • organ that bears no leaves or node • have multiple functions: – 1. anchors the vascular plant in the soil – done by the lateral roots • lateral root – plant organ that functions in increasing anchorage – 2. absorbs minerals and water – mostly done at the tip of the root by root hairs • root hair = thin tubular extension of a root epidermal cell – 3. stores carbohydrates – 4. can undergo vegetative reproduction – 5. production of hormones – e.g. cytokinin and gibberellin (stem growth) • numerous types of roots • but two types of root systems: fibrous and taproot Roots • numerous types of roots – adventitious – form from unusual locations – e.g. leaves or stems – aerial – growth above ground Prop roots. Storage roots. • e.g. orchids – aerating – grow up above the ground or water – coarse – undergo secondary thickening and can be woody – haustorial - seen in parasitic plants; substrate is the body of another plant • roots known as haustoria Aerating “Strangling” aerial roots. Roots • numerous types of roots – prop – exposed adventitious roots produced near the base of the stem – stilt – adventitious roots that grow down from lateral branches off of a stem – storage – for storage of food and water – includes taproots – structural – large roots with secondary thickening, gives support to large woody plants and trees Prop roots. Storage roots. Aerating “Strangling” aerial roots. • found in eudicots and gymnosperms • develops from the embryonic root (known as the radicle) • taproot gives rise to multiple lateral roots Taproot systems – lateral roots can also produce smaller, lateral roots – lateral roots can become swollen like the main taproot • e.g. sweet potatoes and cassava • generally penetrate deeply • are well adapted to deep soils where groundwater is not close to the surface carrot turnip cassava • most monocots – e.g. grasses • mat of generally thin roots that spread out below the soil surface • most of the roots are similarly sized • the embryonic root (radicle) dies early on and doesn’t form a taproot • many small roots emerge from primordial tissues found in the stem • each small root forms multiple lateral roots • does not penetrate the soil deeply • excellent at holding topsoil in place Fibrous Root Systems Root Structure • fairly simple – no leaves, leaf axils, axillary buds etc… • root tip: – tip of the root is where growth in length occurs – root apical meristem (RAM) present in the root tip – Root AM – more orderly than the Shoot AM • there is a quiescent region in the RAM called the quiescent center Root Apical meristem Root cap Root Structure • root cap: – the RAM is protected by a root cap as the root pushes through soil – golgi of the root cap cells secrete a mucilage or slime to help the root push through the soil – cells are small and meristematic Root Apical meristem Root cap • Growth occurs just behind the root tip, in three zones of cells: Root growth – Zone of cell division • region of mitotic division • can also be called the Meristematic zone – Zone of elongation – located just behind the root apical meristem • cells begin differentiation in this region • 1. outermost cells called the protoderm form the epidermis • 2. center cells called provascular tissue form the primary xylem and phloem • as these cells move up and away from the root tip – primary X and P becomes metaxylem and metaphloem • permeable to water Cortex Vascular cylinder Epidermis Root hair Zone of maturation Zone of elongation Apical meristem Root cap Zone of cell division • Growth occurs just behind the root tip, in three zones of cells: – Zone of maturation – also called the root hair zone • vascular bundles contain metaxylem and metaphloem • many of the epidermal cells extend out to form root hairs • innermost cells of the cortex differentiates and form a cylinder called the endodermis – waterproof region that forms around the developing vascular bundle Root growth Cortex Vascular cylinder Epidermis Root hair Zone of maturation Zone of elongation Apical meristem Root cap Zone of cell division Roots: Internal organization • • primary growth of roots produces the epidermis, ground tissue, and vascular tissue from outermost to innermost: – 1. epidermis – from the protoderm of the developing root – 2. cortex – ground tissue – 3. endodermis – encircles the vascular cylinder – 4. vascular cylinder – contains xylem and phloem • also contains a tissue called pericycle – layer of parenchyma or sclerenchyma cells • just inside the endodermis • in dicots – can form lateral roots endodermis Monocot Roots: Vascular Bundle Epidermis Cortex Vascular cylinder Endodermis -Monocot roots: -vascular bundle consists of a core of parenchyma (similar to the pith of a stem) -core is surrounded by a ring of alternating xylem and phloem tissue Pericycle Core of parenchyma cells Xylem Phloem 100 µm Transverse section of a monocot root with parenchyma in the center. The stele of many monocot roots is a vascular cylinder with a core of parenchyma surrounded by a ring of alternating xylem and phloem. Key Dermal Ground Vascular Pericycle Endodermis Xylem Phloem Core of parenchyma cells Dicot Roots: Vascular Bundle Epidermis Cortex Vascular cylinder Endodermis Pericycle -Eudicots and gymnosperms: -vascular bundle is a single vascular cylinder called a stele -surrounded by a pericycle and endodermis Core of parenchyma cells Xylem 100 µm Phloem Transverse section of a typical root. In the roots of typical gymnosperms and eudicots, as well as some monocots, the stele is a vascular cylinder consisting of a lobed core of xylem with phloem between the lobes. Endodermis Pericycle Xylem Phloem 50 µm Key Dermal Ground Vascular LE 35-14 100 µm Emerging lateral root Cortex Vascular cylinder Epidermis Lateral root Lateral roots arise from within the pericycle, the outermost cell layer in the vascular cylinder Plants • two designations: herbaceous (herb – nonwoody) and woody plants • wood: considered to be made of secondary xylem Vascular Cambium • meristematic tissue that forms during primary growth – – – – also called a lateral meristem source of secondary xylem and phloem VC located between the xylem and phloem of the vascular bundle in non-woody plants – the cambium ultimately stops dividing and differentiates into more xylem and phloem Vascular Cambium – BUT in woody plants – the VC never undergoes cell arrest and continue to divide and differentiate • ultimately forms secondary xylem and phloem – made up of two types of cells – fusiform initials and ray initials • fusiform – long and tapered cells that can differentiate into secondary xylem or phloem • ray – shorter and more cuboidal cells that forms more parenchyma cells Secondary vascular tissues • found in woody plants • vascular cambium differentiates into secondary xylem and secondary phloem • secondary phloem – no arrangement of rings or early and late wood • secondary xylem contains all the type of cells that are found in primary xylem – tracheids, vessel elements, sclereid cells and parenchyma Secondary vascular tissues form from secondary growth Cork Cambium • another lateral meristem – forms from the outermost layer of secondary phloem bark (cork) Cork Cambium – cork cambium divides only for a few weeks then differentiates into cork cells (i.e. cork) – cork cambium + cork = periderm tissue • also known as bark – every few years a new CC must form –so several layers of cork and bark build up over the years – the cork and bark are impermeable to water – problem! – specific cork cells round up as they mature and form lenticels secondary phloem LE 35-10 Primary growth in stems Shoot apical meristems (in buds) Epidermis Cortex Primary phloem Primary xylem Vascular cambium Cork cambium Lateral meristems Pith Secondary growth in stems Periderm Cork cambium Pith Cortex Primary phloem Primary xylem Root apical meristems Secondary xylem Secondary phloem Vascular cambium SO what the heck is bark??? – non-technical term – all tissues outside of the vascular cambium – includes the: • secondary phloem • cork cambium • cork (phellum) – may also be called periderm • outer covering of non-woody stems and small woody stems non woody plant? small woody plant? PERIDERM big, honkin’ tree?? BARK Secondary growth: Wood • most commercial dicot woods contain a large amount of vessel elements and fibers – strong and tough (hardwoods) • wood from conifers – fewer vessel elements and fibers and have a softer consistency – softwoods • hardwoods: – – – – maple oak walnut birch • softwoods: – – – – – pine cedar fir spruce redwood Secondary growth: Wood • vascular cambium becomes quiescent during times of stress (extreme hot and cold) and stops forming new cells • when the VC resumes its growth and starts to form new secondary X and P – forms new rings of secondary xylem or wood – first wood that forms – early wood or spring wood – later in the season – late wood or summer wood • late wood + early wood = annual ring • early years of a tree are characterized by vigorous growth – innermost annual rings are wider – more secondary xylem Secondary growth: Wood • oldest regions of the wood found at the center = heartwood (darker) – as the tree grows older the components of the xylem in the inner rings stop conducting water – gradually converted to inactive wood and then heartwood (dead?) • youngest region of the tree is found at the periphery= sapwood – most actively conducting wood Some websites to check out • http://www.slideshare.net/apurvanagvenker/ anatomy-of-dicot-monocot-leaf • http://waynesword.palomar.edu/index.htm • https://www.youtube.com/watch?v=33H93Rlz k2w