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CHAPTER

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CHAPTER OUTLINE
Root Systems

to produce flowers, fruits, and

seeds. Water and minerals, or

Origin and Growth of Primary Tissues

Primary Structure Effect of Secondary Growth on the Primary Body of the Root Origin of Lateral Roots Aerial Roots and Air Roots

Adaptations for Food Storage: Fleshy Roots

inoqganic ions, absorbed by the roots move through the xylem to the aerial parts of the plant. Hormones (particularly cytokinins and gibberellins) synthesized in meristematic regions of the roots are transported upward in the xylem to the aerial parts, where they stimulate gowth and development (see Chapter 27). Roots also synthesize a wide variety of secondary metabolites, such as nicotine, which in tobacco is transported to the leaves (see Chapter 2). In addition, roots function in clonal regeneration (the roots of certain eudicots produce buds that can develop into new shoots), rcdistribution of water witliin the soil (see Chapter 30), and secretion of a rrast arrzy of substances (root exudates) into

-I-h" first structure to emerge from e germinating sed I is the root, enabling the developing seedling to irecome
anchored in the soil and to absorb water. This reflects the two primary functions of roots: anchorage and absorption (Figure 2+l). TWo other functions associated with roots are sorage and conduction. Most roots are important storage organs, and some, such as those of the carrot, sugar bet, and sweet potato, are specifically adapted for the storage of food. Foods manufactured above ground, in photosynthesing portions of the plant, move down through the phloem to the storage tissues of the root. This food may evenrually be used by the root itself, but more often the stored food is digested and the products transporled back through th phloem to the aboveground palts. In biennial plants, such as the sugar beet and carrot" which complete their life cycle over a frweyear perid, laqge fotd reserves accumulate in the storage regions of the root during the fust y:.ar. These food reserves are then used during ttrc second year
558

CHECKPOINTs
After reading this chapter, you should be able to answer the following:

l. 2.
3.

Name the tvro principaltypes of root systems, and describe horl they differ n both origin and tructure-

Wrat changes occur to the rootcap durfng elongatfon of the root, and what are some functions of the rootcap?
What tissues are found in a root at the end of primary growth, and how are they arranged? Describe th ffect of secondary growttr on the primary body of the root

4.
5.

Ii/frele do hteral roots originare and wtry are ey said to be

endogenous?

lnNe 1 a{

i ..i \,4,i +O

t6 /rCi, *\lt

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'r '

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ti
/-

Root

Syfems

559

ttte prinary root, terrned the taproot, grows directly dovrnwan giving rise to branch foots, or lateral roots, Tte older lueral 'roots are found nearer the bas of the roor (wtrcre the root and . . stem rffi), and the younger ones.neane,r the rcot p. This rype :, . l' of root systenHhat is, one formd frbm' a strongly developed primary root and ie b'ranctres*is calteila taproot systsm (Fig. ' ,l .-.1 , ' j,::*ri',Ife

.:

'';,''r-,. l-i.'' ' rln onocots, the primary -"t is usrally shcit-lived, an4 ,' ,i,,'- .idstead, re rnain foot systgp dorelws from t".rs ttrat arisc" ftom '.tib . l j .' stem. Ttrse senr-borne rrx)ts, cornnionly i;alted adpcntitouE '

z.F?A)-

.'

2+l and/r7b,

in nicU no rc loot is more p'rominentlthan dE ors (Ftg

.ures

b,

k)

24-1 Development of root and shoot in a monocot Diagnms

of oat (Avena satva) plants, showing relatire sizes of the root and shoot srstems at (o) 31 dar, (U 45 daF, and (c) 80 days after plantfng. The oat'plant, a monocot, has a fibrou rot srs-tem. The roots are inrrolved primrlly with anchorage and absorption. Each the unts depkted here represents I foot (approximate[. "ertcal 30 centimeters).

the rhizospherg tbe volurne of soil around living plant roots thu is influenced by root activity.

Root Systems
The first root of e plam originares in the ernbryo and is usually callcd the prinary roo In all seed plmts excspd the monocots

Ttre spatial .cunfigruation, or rchitectu$ of a root sy-stem can show considerable wriation everi wiftin different parts of ,a single root systdm. Growing mos are etftrercty sensitirre to a wrde range of enrironnsltal prannters, includipg gravity, ligbt; gradients of moisture, ternperatul, and,nutrients in the soil. A sriking example of tb dorelopmental phsicity, or adapabiliry of tlre rcot system of nuny species is its respmse :to tlre unwen distribution of nimogen and inorganic phoaphate by the preferential and rapid devetopnpnt of lueral rods into nutrient-ricb zres (see Cha$er 29). Ttle extent of a root systnr-at ls, the 'depth to which it penetrates ths sol and the distance it spreads laterallyis dependent on ser'eral factors, including tbe enironmental pararteters just mentioned. Tnproot systems generally penetnate deeper into the soil than fibrous noot systems. The shallowness of fibnous root systems and ttre tenacity with which they cling to soil particles rnake such plants especially well suited as ground cover for the prevention of soil erosion. Most ees have surprisingly shallow root systems, with 90 percent or more of all roots located in the upper 60 centinrcteis (about 2 feet) of soil. The hlk of most fine mots' or so-called feeder noots, the rma actively engaged in the uptake of water and minerals, occurs in the upper 15 centimeters of soil, the part of the soil nonnally richest in nutrients. Many fine roots are heavily infeaed with mycorrhizal fungl (see Chapter l4). Sonrc uroe.s, such as spruces, beeches, ard poplars, rarely produce deep taproots, wtrcreas o{hen, such as oaks and rruny pines, commonly produci relatively deep taproots, making thcse trees difficult to trarisplant. Ttrc recond fo deE 9f pcnetrarion by roots probably be longs 1o Boscia albitrunca (family Caprparidaceae), at 68 meters (over 220 fe), in the central Kalahari Desri:'Roots' of the desen shrub nresquite (Pmsopis juliflora) wer found growing at a depth of 53 mercrs (nearty 175 feet) af an open-pit mine near Tucson, Arizona. During ttrc diggng of the Srez Canal in EglTt, roots of Tizrnarix,and Acacia tnees wre fotnd t r deph of 30 meters. The rcots of alfaHa (Medicago suivtt) may extend to depths .ot'6 rneters or rore. The lateral spread of tee rmts is usually geater-frequently four to seven tinres greatei'-than ttrc spred of the crown of the Eee. The root systms of nraize plmts (Zee mays) often reach a deph of about 1.5 meters, with a lateral spread,of about a nneter on all sdes of th plant One of tk most &failed snrdis on tbe ex[ent of tbe root and shoo systems of any one plant was conducted 'm a forrmonth-old rre plrot (Secale cerealc). Th toal surface rea of the roc systent. inchrding rmt hain. \f,ras 639 square meters,
:

CH A PT

The Root: Strusnre and Development'

2+-2 Taproot and fibrous root

systems Two tlpes of root systetns are represented here by nro'pnirts prlmte. (a)Taproot s)4stem of the blazfnf star (Liatris punctato),a eudicot. fbl Fibrous 'foot system of wire grass (Arlsttdq : . purpuru),a monocot. Each of the vertlcat unlts deplcted here represents

'

''

'

30 centmeters. Taproot srstems . generally penetnte deeper into the soll

than fibrous root systerns.

...

Fine roots are generally short-ve persisting on average for just a few months, alough sorne rnay lire much longer. The shortlived fine roots of ttees are in a state of constant flux, with death and replacenrent ocgrrfuig simultaneously; therefore, the flucnation in the populatin and conoentration of rogts in the soil is as dynamic as that of 'ttrc leaves and twigs in the air above. It has been estimated at,as much as 33 pereent of global annral net produaivity in terrestrial ecosystems is devoted to the production of fine rcots. Even when plants are carefully transplante{ the balance bctween shoot and root is inrariably distued. Most of the ne roots are left behind when the'plant is removed from the soil. Cutting back the shoot helps to reestablish a balance betrreen the root system and the shoot system, as does repotting a pl4qt that is root bound into a larger container.

Thd'llnt'lUaintain a Balancg betweln lts Shogt andRootSystqms:...j..,.'.: : ..,:.::, j. ..,,;;.r :

.

'

The grourth of many rcots

plarit, a balance is. maintained benxeen the totd surface anca arailable for the l rnanufacture of food: (the photosynthesinng surface) and he surface Err.:available for the absorption of wdcr and minerals. This functionat balance, which is between the fine mots and leaf arrlq,csn.bc expressed as a ratio, the root:shoot'ratio. In seedlings, the total water- and mineral-absorbing surface usually far exceeds the photosynthesizing surface. Hcqever, the root:sht, atio.decreases gndu-

In a

a cgntingoru proces$ ttn!. stops onlf unde sqc adverse conditionq as {ougtn md fow !em-, peratrf{E$. Durlng their grg\r thr-oogh rs so4, rgqts follm, the pa of least resistance and fteqently follow spaces left by wlier rmts that have died and rotted.

Origin qni,Groiilth 9{

Brimaryli:ues

,, ,

n Oppg*Rtly

The Tip of the Root ls (overed by a Rootcap,

WhichProducesMucilage,]',-,..,-,,i,,.,
The motcap is a thimblelike nrass of living paredchyma cells that proteca the apical rneristem behind it and aids the root in

ally
ing

as a plant ?gGS.,

its penetration of the soil (Figures 2+3 thogh

2)-

As ttle

ff damagg

to tbc rod systfm saimsly reces its absorb

grqnrth is reced by lack of

sufu, shot

wm, mitsals.

root gross longer and the roo{cry is pushed foflrar alls on tbc peripbry of the rootry,rcctrEte largp arxrnts sf nudryc

0rigin and Growth of Primary Tissues

561

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Root hain
MucLagre

sheath

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: (b)

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24-4 Mucilage sheath on root tip fol Mucilage

o.n thB rootcap of a rnaize (Zeo moys) root, containng border cells. fb) Dark-field image of a living root showing a "cloud'of border cells suspended

inthemucilagesheath,whichlsnotapparentin.thlsvlew.
. .,':t)

24-3 Eudicot root A portion of a eudlcot rool showing the

spatial relatlonshlp between the rogtcap and the regon of root halrq, and (near the top) the sltes 9f emergence of the latenl roots, whlch arlse from deep wlrln the parent rtoot New root halrs arlse

just behind the reglon of elongatlon at about the same rate as the older hairs die off. The root tip ls,covered by a.rnucilage sheath, whlch lubrlcates the root durlng its passage through the soll.

(a highly hydrated polysaccharide), which lubricates the root during'itS phssage through the soil (Figure 2+4). Eventually, these periphal cells are :released from the rootcap. Called border cctk, these rootcap cells are programrned to separate from e rmcap md from eac other as t@ reac e rootcap periphery- Oo their release- ttre bmder cells-which may

remain' llirc in the rhizosphere for several weeks--undergo changes,'in gene expression that enable them to iiioduce and exude Specinc proteins thar are completely differenr'fioni those of thb rootcap. As the border cells are release 'new'cells are added to'the'r)tcap. The number of border cells relerised each day varies, in part" with the plant farnily. For example, as fs' as l0 border cells are shed daily for tobacco (family Solanaceae), compared with as many as 10.000 for cotton (farnily Malvaceae). Border cells and their products may contribute up to 98 percent of the weight of carbon-rich material released into the sol as root exudates. with rising levels of saou dioxide in ttp atmosphere and associated climue change, the role of soil in sequestering carbon is of grear interest. Sewral fimctions harrc treen anributed to bsder cells md ttgir exsddes. Among these functions are protwrion of e arcal