LSM 3261 Life Form and Function

Roots and Mineral Nutrition

Lecture 4

Reference Definition of the root Main functions Root types

Taproot Fibrous root

Lecture 4 Topics

Root structure
Tissues Root R t cap Root hairs Herbaceous eudicot or monocot root structure Cortex (including endodermis) Stele Water movement in the root Vascular cambium Periderm Lateral root development

Root modifications

Reference
Solomon, E.P., L.R. Berg and D.W. Martin. 2011. Biology. 9th ed.
Chapter 36

Definition of the Root

A root is typically a subterranean or aquatic multicellular axis or q appendage of a plant the function of which is the absorption of mineral salts in solution and anchorage of the plant. (There are exceptions to this.)

Water hyacinth, with roots growing into water. Taeniophyllum obtusum, a shootless, leafless orchid

Main Functions
Anchor plants Absorb water and nutrient materials from the soil Conduct water and nutrient materials to the rest of the plant Storage Serve as food reserves for the plant (roots specialized p ( p for even more storage are covered later under root modifications)

Root Types
Taproot Fibrous root

 Taproot = primary root In gymnosperms and eudicots Forms from the enlarged seedling radicle Radicle is the embryonic or primary root Travels directly down into the soil Usually travel deeply into the soil to capture water t Collects water and nutrients from deep in soil

Taproot 1

 Most large trees lose their taproot and develop a shallow root system that sends branches deep into the soil Secondary root = those which d h hi h directly arise from the tap root Tertiary root = those which directly arise from the secondary root

Taproot 2
roots trunk k

primary root secondary root

Fallen casuarina tree (Casuarina equisetifolia)

tertiary root
Tongkat Ali (Eurycoma longifolia)

 In monocots Primary root dies quickly Roots develop from the stem or leaves called adventitious roots (adventitious organs are those that arise in unexpected locations)

Fibrous Root 1

 Cling strongly to the soil prevent surface soil erosion Collects surface water and nutrients Secondary root = those which directly arise from the fibrous root Tertiary root = those which directly y y arise from the secondary root

Fibrous Root 2

secondary root

fibrous root

tertiary root

Root Tissues
Mature tissues M i Dermal system Epidermis p Root cap Root hairs P id Periderm Ground/fundamental system Cortex Stele Vascular system Stele Primary vascular tissues Secondary vascular tissues Meristematic tissues Apical meristem Vascular cambium Cork cambium Anomalous forms (not covered in this module)

Root Cap
Root apical meristem (area of cell division)

Root cap

Hollow conical ll i l layer of multiple layers of tough cells Cells are sloughed off as the root forces its way through soil; replaced by cells the divide and grow inside th i id the root cap t May aid in geotropism removal disorients the apical meristem i l it

250 m

Root Hairs

Tubular extensions of single epidermal cells No cuticle so absorptive Behind the growing tip A modified trichome much longer than most trichomes yet i h still relatively short maximally ~1 cm Very numerous Have very close contact with water/soil particles Increase the surface area of the root Increase the absorptive capacity of the root

Herbaceous Eudicot and Monocot Roots

Epidermis Cortex

Stele

Eudicot Root

250 m

Roots in a primary state of growth, i.e., i e those of herbaceous eudicot or monocot Have three basic layers of tissues/zones Epidermis Cortex Stele

Monocot Roots
Stele S Epidermis cell Cortex cell

Pericycle cell Pith cell Xylem vessel element Phloem cell

Monocot roots are variable in internal structure Usually with a pith in the center All monocots have no vascular cambium but may have other forms of secondary growth, e.g., periderm e g periderm, anomalous forms (not covered in this module) )

Monocot Root

250 m

 Cortex = Tissues immediately below the epidermis up to y p p and including the endodermis Parenchyma cells (storage) Sclerenchyma (support in persistent roots) Collenchyma (rare in roots) Innermost layer of tightly packed endodermis connected together by casparian strips, which are rich in waxy suberin

Cortex (including Endodermis)

Pandan (Pandanus sp.) has persistent roots

Stele
Stele = Central cylinder of tissues consisting of
Cortex cells filled with amyloplasts Endodermis cell Pericycle cell Phloem cell Xylem vessel elements Intercellular space p 25 m

P i l = Si l layer Pericycle Single l of parenchyma cells inside the endodermis which separates the vascular tissues from the endodermis Vascular tissues = Xylem a d p oe and phloem in alternating a e a g and radial patches Intervascular regions = Ground tissues of G d i f parenchyma and/or sclerenchyma between the xylem and phloem patches

Stele
Stele S

Stele (continued)
Pith = Centermost tissue ground Epidermis consisting of gro nd cell tissues of parenchyma and/or sclerenchyma and y Cortex cell usually absent in dicot roots
Pericycle cell Pith cell Xylem vessel element Phloem cell

Monocot Root

250 m

Announcements and Discussion

Practical 2 on Roots and Reproductive Structures
2-4 pm at LS Lab 7 on Thursday, 8 Sep 2011 Come punctually for the detailed pre-lab briefing Bring the usual items as for Practical 1

Tutorial 2 on a structure to function experiment

Download it after 5 Sep

Appeal for help with field work

Assist my three Honours students with their final year projects Abandoned rubber plantations Wasteland vegetation A t l Autecology of potentially invasive tree f t ti ll i i t Chance to see what ecological research involves Good experience for field-oriented/adventurous students field oriented/adventurous

Water Movement in the Root 1

Water and minerals move into root by four p pathways: y
Apoplastic: via porous connected cell walls extending from the root hairs S Symplastic: via plasmodesmata i l i i l d interconnected d cytoplasm of adjacent cells Transmembrane: via successive crossing of plasma membranes and cell walls of the cells along the p pathway ( y (not in main text) ) Combination of the above pathways (not in main text) http://www.youtube.com/watch?v=o32jqyIpoHg

Water Movement in the Root 2

The casparian strip of each endodermal cell blocks the passage of water and nutrient minerals past the cell walls between the endodermal cells and directs water into b h d d l ll d di i endodermal cytoplasm to the phloem or xylem from the apoplast Xylem conducts water and dissolved nutrient minerals from the root hairs Phloem conducts sugars from the leaves as well as sugar from stored nutrients in the root cortex

ftp://tdata.atu.edu/cindy.jacobs/Intro%20to%20Biology/Biol%201014%20Unit%202%20Spr%2007/Ch%2018/animations/root_functioning.html

Water Movement in the Root 3

Root hair/ground epidermal cell cortex including endodermis pericycle root xylem

N.B. N B Transmembrane or combination pathways not illustrated here

Learning Tip: Using Analogies

Analogy = A likeness between two entities, not based on the th resemblance of the two things, but of a few of the bl f th t thi b t f f f th key attributes of those things Th more key attributes in common, the better The k tt ib t i th b tt Helps in understanding new concepts or ideas, but has limitations Example
Analogy for a stoma
Hole in a wall Window in a wall Automated window in a wall

Not all analogies are equally effective; some are better than others

Vascular Cambium 1
Found in most older eudicot and all older gymnosperm roots Develops in the parenchyma of the intervascular region between the primary p y xylem and primary phloem (cell divisions start) t t) When cell division extends to the pericycle, the vascular cambium is fully developed

Precursor of vascular cambium

Vascular Cambium 2
Vascular cambium divides to form xylem inside, phloem outside
Primary phloem pushed outward Primary xylem stays y y y where it is

Ring of vascular cambium becomes circular by extra divisions of vascular cambium between the xylem arms
Primary phloem, epidermis and cortex are crushed and lost as outer bark Periderm replaces epidermis with increase in diameter of ith i i di t f root

Periderm 1
Can originate from the cortical parenchyma cells just below the epidermis to the pericycle Like the periderm in the stem

Precursor of vascular cambium

Periderm 2
The source tissue divides to form the cork cambium to form columns of cells:
Those outwards become cork cells or phellem which are dead a maturity and have a mechanical protection and h i l i d insulating functions Those inwards become phelloderm or cork parenchyma and have a storage or cork cambium reserve f function. (When growth i (Wh h stops as during a the dry season or winter, cells differentiate into i t cork or phelloderm. When k h ll d Wh growth resumes, the phelloderm cells start dividing and become the cork cambium.) cambium )

Periderm 3
Several sequent periderms form the bulk of the bark which is all the tissues outside the vascular cambium
cork
cork cambium

cork
cork cambium

cork
cork cambium

cork
cork cambium

cork parenchyma

cork parenchyma

cork parenchyma

cork parenchyma

cork
cork cambium

cork
cork cambium

cork
cork cambium

cork parenchyma

cork parenchyma

cork parenchyma

cork
cork cambium

cork
cork cambium

cork parenchyma

cork parenchyma p y

cork
cork cambium

cork parenchyma

Lateral Root Development D l

Originate in the pericycle Cells in one portion divide and pushes through the endodermis, the rest of the cortex and epidermis Each lateral root has all the parts of the larger root from which it originates
Root cap Root hairs Epidermis p Cortex (including endodermis) Stele (with connections to the main root developing)

Ruptured epidermis and cortex Lateral root Cortex

100 m

Root Modifications
Root modifications include Aerial roots B ttress roots Buttress Contractile roots Haustorial roots i l Pneumatophores Prop roots g Storage roots
o Food o Water

Digression: Alternative Classifications

Classification = Hierachical set of groups and subgroups based on certain criteria Same set of entities will produce different classifications based on different criteria Classification of roots Based on origin Primary (tap), adventitious (including fibrous), secondary, tertiary Based on modifications or not Aerial, buttress, haustorial, pneumatophore, prop, storage, storage normal ( unmodified) (= nmodified) Using categories from one classification to fit into those of another usually does not work so be aware of which classification is being used at any one time

Aerial Roots
Aerial roots =
Adventitious roots which Ad ii hi h arise in the aerial parts of the stem for clasping supports or not
Aerial roots

Phalaenopsis hybrid

Money plant (Epipremnum aureum)

Why are aerial roots y considered adventitious?

Buttress Roots 1
Buttress roots = B tt t
Roots with a board-like or planklike lik growth on their upper sides, th th i id presumably for a supporting function
Which roots are buttresses outgrowths of?

Buttress Roots 2
What Wh are the functions of h f i f buttress roots?
gravity pulls the t th tree downwards compression strut

tree tr runk

tension strut t t

20 m

Contractile Roots
Contractile roots =
Roots often associated with corms or bulbs which shorten by becoming broader to pull the corm or bulb deeper into the soil (e.g., gladiolus (Gladiolus sp.)) Why does the contractile root broaden when it becomes shorter? (Hint: The volume of the cell stays the same.)
Contractile roots

Haustorial Roots
Haustorial roots =
Roots of parasitic species which R f ii i hi h grow into the host

Mistletoe (Macrosolen cochinchinensis) )

Pneumatophores 2
Pneumatophores =
Breathing roots in plants growing in marshes or swamps

Api api (Avicennia sp.)

Do pneumatophores branch off from secondary roots?

Prop roots =
Adventitious roots which arise from the lower nodes of the stem which have a supporting function

Prop Roots

Maize (Zea mays)

1.93 m

Bakau (Rhizophora sp.) ( p p)

Food Storage Roots

Storage roots =
Roots modified to have much storage parenchyma containing starch or other products d t Chinese radish

Sweet potato

Food Storage Roots

Storage roots =
Roots modified to have much storage parenchyma containing starch or other products d t
Left to right: carrot (Daucus carota), sweet potato (Ipomoea batatas), Chinese radish (R h di h (Raphanus sativus i Longipinnatus, bangkwang or yam bean (Pachyrrhizus erosus), tapioca (Manihot esculenta)

Record-Breaking Carrots

John Evans grew the Worlds heaviest World s carrot, at 8.61 kg in 1998 in Alaska, USA.

The world s longest carrot was 5.841 m long and world's 5 841 grown by Joe Atherton (UK). The carrot was measured at the UK National Giant Vegetable Championship in Somerset UK on 2 September Somerset, UK, 2007.

Water Storage Roots

Desert rose (Adenium obesum)

Water storage stems and roots Will see this sp. during Botanic Gardens fi ld trip G d field t i Auspicious plant of the Chinese