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Roots Botany 313 Plant Structure Tamra Prior Overview: • • • • • • • • • • Evolutionary Origins Root Growth & Development Structures & Tissues Monocots vs. Dicots Root Functions Apoplastic and Symplastic Transport Types of Roots Nitrogen Fixation Mycorrhizae Weird Plants that are exceptions Root Origins • The first vascular plants had rhizoids that absorbed water and nutrients. • Rhizoids are delicate uni- or multicellular root- and hair-like structures that function in anchorage and absorption but lack xylem and phloem. • Roots evolved in response to the pressures of a land environment and increasing plant size. • Roots have changed relatively little over time, likely because of their protective subterranean environment, they were not exposed to the same selective pressures as the stem. Root Origins • Seed plant roots may be considered to be: 1.) A modified stem or 2.) An entirely new organ that evolved independently of the stem. • The first view is accepted now because the roots of extant plants are very similar to stems of ancestral fossil plants. • Roots of most plants are protostelic (like the stems of very primitive plants); roots with a central pith have an alternate arrangement of xylem and phloem that may reflect a protostelic origin. Radicle To Root ©1996 Norton Presentation Maker, W. W. Norton & Company Radicle becomes the Root What about here? Root Tip Pericycle and Lateral Root Growth • Here you can see two root apices initiating from the pericycle. • What is the pericycle doing? Growing out through cortex Root cap Zone of cell division • In monocot plants, there are few lateral root formations. Monocots rely on many new fibrous roots • Notice their connection to the ridges of xylem Maturation of Dicot Root Dicot Vascular Cambium Maturation Shape Monocot Root Root Vascular Cylinder and Cortex Ranunculus acris - buttercup Cortex Endodermis Phloem Meta-xylem Pericycle Endarch: protoxylem is inside the metaxylem protoxylem is outside √Exarch: the metaxylem Proto-xylem Monocot vs. Dicot Root Functions 1. Roots anchor plants to soil; provide stability to above ground parts of the plant. 2. Roots absorb water and dissolved mineral nutrients from soil and transport them to the stem. cortex cell epidermal cell root hair penetrates soil spaces ©1996 Norton Presentation Maker, W. W. Norton & Company Root Hairs intercellular gas space + to vascular cylinder voids with air space water Root Functions 3. Roots can be used for storage of food reserves. 4. Roots can determine the longevity of the plant – annual, biennial or perennial. In biennial plants, large food reserves accumulate during the first year of vegetative growth. Then they are used to produce flowers, fruits and seeds the second year. In perennial plants, the food reserves allow the root to survive adverse environmental conditions and then produce new vegetative growth in the spring. Do roots of annuals store a lot of food? Why or why not? 5. Roots are sites of hormone production – especially cytokinins and gibberellins, which are then transported upward in xylem to stems and leaves. Endodermis Xylem inside • The endodermis is thus responsible for selective mineral uptake. Suberin- waxy barrier to apoplastic movement Cortex outside • minerals cannot go between cells • minerals must go through cells • Cell membrane proteins (active transporters) determine which minerals may be taken up Close-up of Transport through Endodermis Root Types • Tap Root- The primary root of a plant formed from the radicle of the embryo; a stout tapering main root from which arise smaller lateral roots or branches. • If the radicle or taproot is damaged, a lateral root emerges and assumes the dominating role. Growth of the taproot continues throughout the life of the plant. This is common in gymnosperms and dicots. • Lateral or Branch Roots- A root that arises from another older root, arises from the pericycle. Root Types • Fibrous Roots – A root system in which no one root is more prominent than the others – all roots are more or less the same size (at least in diameter – length may vary), typically the plant has a mass of these. • Common in monocots • Arise because the radicle dies during or soon after germination; usually adventitious in origin. • Plants may be perennial or annual. While individual roots are small, the mass of them have enough stored food reserves to allow the plant to survive more than one growing season. Root Types • Adventitious Roots- Roots that develop from organs other than pre-existing roots (so leaves and stems!) • The root system of grasses is all or mostly adventitious. Also found in ferns, clubmosses and horsetails. • Contractile Roots- Roots that contract to pull the shoot, corm, or bulb down deeper into the soil. Roots extend through the soil and become firmly anchored. The uppermost parts begin to contract and the stem, etc. is pulled downward so it buries deeper. • This is caused by changes in the shape of cortical cellsthey expand radially and shorten, losing ½ - ⅔ of their height; vascular tissue buckles but does not lose its function. Narcissus Root Types • Aerial Roots- Are adventitious roots produced from above ground structures; in ivy (Hedera), the roots cling to the surface of objects (trees or walls) to support the climbing stem. • Velamen- A multiple layered epidermis that surrounds the aerial roots of some epiphytic orchids and provides mechanical protection for cortex and reduction of water loss. There are many specialized types of aerial roots: • Prop Roots- Roots that serve for support, as in corn. They can branch down from lower nodes of the stem or drop down from branches as in some trees. They can enter the soil and absorb water and nutrients. Examples: Mangroves, Banyan tree, palms. Root Types • Pneumatophores- Roots that grow upward from submerged roots in mud/water; found in trees that grow in swamps, such as mangroves • Do help to anchor the plant, but main function is to aerate the plant; allows air to travel through the root system and provide O2 for respiration; roots have aerenchyma and lenticels. • Buttress Roots- In shallowly rooted trees at the base of the trunks. • They may be up to 4 meters tall; and are specialized for support with large amounts of fibers and small amounts of xylem. • Where else have you heard the word buttress before? Root Types • Storage Roots- most roots store food reserves, some plants have roots specialized for this function. • They are usually fleshy because of abundance of storage parenchyma permeated by vascular tissue; the upper portion of most fleshy roots actually develop from the hypocotyl. • Examples: sweet potato, sugar beet; roots of some desert plants (they can store more than 70 kg of water!) Haustoria (Haustorium) • Haustoria- specialized roots on plants that are parasitic; they are modified structures that penetrate the host tissue to transfer nutrients from the host to the parasite • A primary haustorium is a direct outgrowth of the radicle • A secondary haustorium is a lateral organ that develops from a modified adventitious root or from outgrowths of roots or stems • Examples: mistletoe (Viscum), dodder (Cuscuta), broomrapes (Orobanche) • In some species, the haustorium penetrates the host tissue to the xylem and forms a continuous xylem bridge. • In others, the haustorium forms a mantle of parenchymatous tissue around the host organ. This induces the host to form a placenta-like outgrowth of vascular tissue to supply nutrients to the parasite. Rhizosphere Region of space around the whole root system including the soil between roots of the same plant. Nitrogen Fixation • Root Nodules- enlargements found on the roots of many plants inhabited by nitrogen-fixing bacteria. • Plants cannot use atmospheric N2, but some bacteria can convert it to a form plants can use. • The bacteria provide the enzymes needed for nitrogen fixation; the host plant provides the sugar to run the operation. • Examples” Fabaceae (pea family) and the bacteria Rhizobium; Alnus (alder), Ceanothus (tea bush), Myrica (sweet gale) and Elaeagnus (oleaster) are usually infected by an actinomycete Frankia. Cycads have root nodules containing cyanobacteria – Nostoc or Anabaena; the water fern Azolla may also be infected with Aanabaena. • Infestation of the host root occurs through root hairs. • The plant starts forming nodules in response and bacteria aggregate within cells in the center of the nodule. • Bacteria are surrounded by parenchyma tissue with vascular bundles and an endodermis. • The endodermis retards entry of O2 to the nodule, which would adversely affect N2 fixation. • Plasmodesmata connect the bacteria-containing cells with surrounding parenchyma cells. Mycorrhizae • The surface area of root tips and root hairs are relatively small. • In times of drought, the volume of soil that provides water and minerals can be depleted. • Mycorrhizae greatly enhance the roots’ absorptive capacity. • This is a symbiotic relationship between a plant and a fungus. • The fungus can be either an ascomycete, an basidiomycete or an zygomycete. • 90% of all terrestrial plants are mycorrhizal. • Mycorrhizae have NOT been found in Brassicaceae, Cyperaceae, and Cucurbitaceae. • Ectomycorrhizae- the fungus forms a dense sheath of hyphae around the root tip. • Hyphae extend from the sheath into the soil and also penetrate the epidermis and cortex of the root. • The Hartig net is a much branched network of hyphae in the intercellular channels of the parenchyma cells. These cells function as transfer cells, transporting photosynthates to the fungus and water and minerals from the fungus to the plant. • Endomycorrhizae- the most common type (80% of vascular plants) • The fungal hyphae do not form a sheath around the root; the hyphae penetrates the roots and grows within the cortex. Hyphae penetrate cells walls but not the plasma membranes. • Complex much-branched structures called arbuscules are formed by the hyphae that are enclosed by plasma membranes (but not within the protoplast). • This is an efficient region of transfer of photosynthates to the fungus, as well as water and minerals to the plant. This is also called vesicular-arbuscular mycorrhize. Cool Exceptions to the Rules! This is Dendrophylax lindenii , and the green things are photosynthetic roots (chlorophyll in its root cortex). The leaves are microscopic or non-existent. • A unique (and really cool) aerial root is found in the epiphyte Dischidia rafflesiana (Asclepiadaceae), a.k.a. the flower pot plant. • Some leaves form hollow containers that collect debris and rainwater. Ant colonies live in the “pots” and add N2. • Roots form at the node above the leaf and grow downward into the pot to absorb water and nutrients. Look a monocot with a tap root! Cordyline australis Do you have questions? Do you need to see a previous slide?