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we'll discuss the transition from vegetative shoot formation to reproductive shoot formation.

In describing plant organs, we'll draw examples mainly The basic morphology of most vascular plants reflects their evofrom the two major groups of angiosperms: monocots and lutionary history as terrestrial organisms that inhabit and draw eudicots (see Figure 30.13). AP Biology Reading Guide Chapter 35: Plant Structure, Growth, and Development

The Three Basic Plant Organs: Roots, Stems, and Leaves

Roots and above ground. They must absorb water and minerals from below the ground surface and and light from above the A root is a multicellular organ that anchors a vascular plant in ground surface. The ability to acquire these resources arose from Name ____Jyothis James__________ Period ____1_____ the soil, absorbs minerals and water, and often stores carbothe eo.-olution of three basic organs-roots, stems, and leaves. hydrates. Most eudicots and gymnosperms have a taproot sysThese35: Plant Structure, and a shoot system. evelopment consisting of one main vertical root, the taproot, that Chapter organs form a root system Growth, and D the latter tem, consisting of stems and leaves (Figure 35.2). With few excepdevelops from an embryonic root. The taproot gives rise to tions, angiosperms and other vaseuIar plants rely rompletel:y on lateral roots, also called Concept 35.1 The plant body has a hierarchy of organs, tissues, and cells branch roots (see Figure 35.2). in both systems for survival. Roots are typically nonphotosynthetic many angiosperms, the taproot stores sugars and starches and stan-e unless photosylldrates. the sugars and other carbohythat the plant will consume during flowering and fruit pro1. This concept is organized into three sectionsplant organs, tissues, and cells. Begin by drates produced during photosynthesis, are imported from the duction. For this reason, root crops such as carrots, turnips, shootdefining a tissue and an organ. on the wasystem. Conversely, the shoot system depends and beets are harvested before they flower. Taproot systems ter and minerals that roots absorb from the soil generally penetrate deeply and are therefore well adapted to Vegetative growth-production of nonreproductive leaves. A tissue is a group of cells with a common function, structure or both. groundwater consists of the surface. deep soils where the An organ is not close to several stems, and roots-is only one stage in a plant's life. articular types of tissues that together carry out pManyplants functions. In seedless vascular plants and in most monocots, such as
also undergo reproductive growth. In angiosperms, reproduc grasses, the embryonic root dies and does not give rise to a tive shoots bear flowers, which are composed oflea\'eS that are main root. Instead, many small 2. The three plant organs are ___roots____, ___stems___, and __leaves__ . roots grow from the stem. Such highly modified for sexual reproduction. Later in this chapter, rootsare said to beadvelltitiOUS (from the Latin advelltiCUS, ex-

resources from two very different environments-below ground

3.

On Figure 35.2, label the shoot system, root system, apical bud, axillary bud, and root system.
traneous), a term describing a plant organ that grows in an unusual location, such as roots arising from stems or lea\'eS. Each small root forms its own lateral roots. The result is a fibrous root system-a mat of generally thin roots spreading oul below the soil surface, with no root functioning as the main one (see Figure 30.13). Fibrous root systems usually do not penetrate deeply and are therefore best adapted to shallow soils or regions where rainfall is light and does not moisten the soil much below the surface layer. Most grasses have shallow roots, con centrated in the upper few centimeters of the soil. Because these shallow roots hold the topsoil in place, grass makes excellent ground cover for preventing erosion. Although the entire root system helps anchor a plant, in most plants the absorption of water and minerals occurs primarily near the tips of roots, where vast numbers of tiny root hairs increase the surface area of the rool enormously (Figure 35.3).
<II Figure 35.3 Root hairs of a radish seedling. Root halTS grt;MI by the thousands Just behind the tip of each root. By Increasing the surface area, they greatly enharn:e the absorption of water and mlneraJs from the SOlI.

Reproduct....e shoot (flower) Apical budl-----,---..Ij

Apical---'t

b,'

shoot

----'llt.

Shoot system

Leaf Axillary

b,'

",m--::::;;;;o;;;;;OOiiiiiiiiii

Root

system

4. Figure 35.2 root and then flowering plant. The plant Define An overview of a explain the difference between a taproot and lateral roots. body IS dMded Into a root S)'5tem and a shoot system, connected by vascular IlSSUe (purple strands In thIS diagram) that IS contmuous Root - A root is a mThe plant shown isorgan that anchors a vascular plant in the soil, absorbs minerals and throughout the plant ulticellular an idealized eudicot. water, and often stores carbohydrates. Taproot and Lateral root - Most eudicots and gymnosperms have a taproot sys- tem, consisting of one main vertical root, the taproot, that develops from an embryonic root. The taproot gives rise to lateral roots, also called branch roots. 5. This photograph shows the root hairs of a radish. What is the function of root hairs?
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Plant Structure, Gro...1h, and Development

739

Although the entire root system helps anchor a plant, in most plants the absorption of water and minerals occurs primarily near the tips of roots, where vast numbers of tiny root hairs in- crease the surface area of the root enormously

6.

What is the advantage of apical dominance to a plant?

By concentrating resources on elongation, the evolutionary adaptation of apical dominance increases the plant's exposure to light. 7. 8. The main function of a leaf is __to perform photosynthesis__. What are five additional functions that modified leaves can perform? a. Tendrils - The tendrils provide support by clinging on to other objects b. Spines provide protection for the plant c. Storage leaves specialize in storing water d. Reproductive leaves produce plantlets which fall off and reproduce e. Bracts- their bright colors attract pollinators Plants have three types of tissues. Place the name of each tissue type and its function in the table below. a. dermal tissue system - is the plant's outer protective covering. In addition to protecting the plant from water loss and disease, the epidermis has specialized characteristics in each organ. They can also provide defense against in- sects by forming a barrier or by secreting sticky fluids and toxic compounds vascular tissue system - carries out long-distance trans- port of materials between the root and shoot systems. The h...o types of vascular tissues are xylem and phloem. ground tissue system - Ground tissue that is internal to the vas- cular tissue is known as pith, and ground tissue that is external to the vascular tissue is called cortex. The ground tissue system is not just filler. It includes various cells specialized for functions such as storage, photosynthesis, and support.

9.

b.

c.

10. What is the function of the cuticle? a waxy coating on the epidermal surface, helps prevent water loss. 11. Xylem conducts __water and dissolved minerals upward from roots into the shoots__. 12. Xylem transport tends to be in one direction, but phloem transport is more complicated. Explain the pattern of sugar flow in phloem tissue. Phloem transports sugars, the products of photosynthesis, from where they are made (usually the leaves) to where they are needed-usually roots and sites of growth, such as developing leaves and fruits. 13. The two major tissues of the ground tissue system are pith and cortex. Where are they found in the plant? Tissues that are neither dermal nor vascular are part of the ground tissue system. They are found between the dermal and vascular tissue. 14. a a. Parenchyma cells - perform most of the metabolic functions of the plant, synthesizing and storing various organic products. Most parenchyma cells retain the

ability to divide and differentiate into other types of plant cells under particular conditions-during wound repair, for example. b. Collenchyma cells - Grouped in strands or cylinders, collenchyma cells help support young parts of the plant shoot. At maturity, collenchyma cells are living and flexible, elongating with the stems and leaves they support c. Sclerynchyma cells - Also functioning as supporting elements in the plant, but with thick secondary walls that are usually strengthened by lignin, sclerenchyma cells are much more rigid than collenchyma cells. Mature sclerenchyma cells cannot elongate, and they occur in regions of the plant that have stopped growing in length. Sclerenchyma cells are so specialized for support that many are dead at functional maturity, but they produce secondary walls before the protoplast (the living part of the cell) dies. d. Xylem cells- Tracheids are long. thin cells with tapered ends. Water moves from cell to cell mainly through the pits, where it does not to cross thick secondary walls. Vessel elements are generally wider, shorter, thinner walled, and less tapered than the tracheids. They are aligned end to end, forming long micropipes known as Water-Conducting Cells of the Xylem vessels.
The two types of water-conducting cells, tracheids and ,"essel elements, are tubular, elongated cells that are dead at functional maturity. Tracheids are found in the xylem of nearly all vascular plants. In addition to tracheids, most angiosperms, as .....ell as a few gymnospenns and a few seedless vascular plants. have vessel elements. \'<'hen the living cellular contents of a tracheid or vessel. element disintegrate, the ceIJ's thickened walls remain behind. forming a nonliving conduit through "''hich water can flow. The secondary walls of tracheids and vessel dements are often interrupted by pits, thinner regions where only primary walls are present (see Figure 628 to review primary and secondary waIls). Water can migrate laterally between neighboring cells through pits. Tracheids are long. thin cells with tapered ends. Water moves from cell to cell mainly through the pits, where it does not to cross thick
secondary walls. The secondary walls of tracheids are hardened with lignin.....itich prevents collapse under the tensions ofwater transport and Tracheids and vessels
I

100l!m

also provides support. Vessel elements are generally wider, shorter, thinner walled, and less tapered than the tracheids. They are aligned end to end, forming long micropipes known as 'essels. The end walls of the vessel elements have perforation plates that enable water to flow freely through the vessels.

(co'onzed SEM)

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Periorat

i. e. Phloem - Unlike the water-conducting cells of the xylem, the sugar- conducting cells Sugar-Conducting phloem are aPhloem unctional maturity. In seedless vascular plants and of the Cells of the live at f Unlike the water-conducting cells of the xylem, the sugargymnospenns, sugars and other Sieve-tube elements: organic nutrients are transported through long, longitudinal view (lM) conducting cells of the phloem are alive at functional maturity. narrow cells Gilled sieve cells. In the phloem of angiosperms, these nutrients are In seedless vascular plants and gymnospenns, sugars and other transported through sieve tubes, which consist of chains of cells called sieve tube or organic nutrients are transported through long, narrow cells sieve tube members. Gilled sieve cells. In the phloem of angiosperms, these nutrients
are tnmsported through sieve tubes, which consist of chains of cells called sieve-tube or sieve-tube members. Though alive, sieve-tube elements lack a nucleus, ribosomes, a Sieve-tube element (left) distinct vacuole, and eytoskeletal elements. This reduction in cell and companion cell: contents enables nutrients to pass more easily through the celL cross section (TEM) The end walls between sieve-tube elements, called sieve plates, haw pores that facilitate the flow of fluid from cell to cell along the sieve tube. Alongside each sieve-tube element is a nonconducting cell called a companion which is connected to the sieve-tube element by numerous channels, plasmodesmata (see Figure6.28). The nucleus and ribosomes of the companion cell setW not only that cell itself but also the adjacent sieve-tube ekment In.some plants, companion cells in leaves also help load sugars into the sieve-tube elements, which then transport the sugars to other Sieve plate Companion cells

Vessel elements. with perforated end walls

\ f---fl---

elements Plasmodesma

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Sieve-tube elements: longitudinal view
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Sieve plate with pores (SEM)


Plant Structure, Gro....1 h, and Development

745

tapered than the tracheids. They are aligned end to end, forming long micropipes known as 'essels. The end walls of the vessel elements have perforation plates that enable water to flow freely through the vessels. Vessel elements. with perforated end walls

Sugar-Conducting Cells of the Phloem


Unlike the water-conducting cells of the xylem, the sugarconducting cells of the phloem are alive at functional maturity. In seedless vascular plants and gymnospenns, sugars and other organic nutrients are transported through long, narrow cells Gilled sieve cells. In the phloem of angiosperms, these nutrients are tnmsported through sieve tubes, which consist of chains of cells called sieve-tube or sieve-tube members. Though alive, sieve-tube elements lack a nucleus, ribosomes, a Sieve-tube element (left) distinct vacuole, and eytoskeletal elements. This reduction in cell and companion cell: contents enables nutrients to pass more easily through the celL cross section (TEM) The end walls between sieve-tube elements, called sieve plates, haw pores that facilitate the flow of fluid from cell to cell along the sieve tube. Alongside each sieve-tube element is a nonconducting cell called a companion which is connected to the sieve-tube element by numerous channels, plasmodesmata (see Figure6.28). The nucleus and ribosomes of the companion cell setW not only that cell itself but also the adjacent sieve-tube ekment In.some plants, companion cells in leaves also help load sugars into the sieve-tube elements, which then transport the sugars to other Sieve-tube elements: longitudinal view (lM)

Sieve plate Companion cells

\ f---fl---

elements Plasmodesma

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Sieve-tube elements: longitudinal view

i. 15. Compare and contrast the following Structure, Gro....1h, and Development 745 structures: CHAH(1 THIUYflV( Plant a. Tracheids are long. thin cells with tapered ends. Water moves from cell to cell mainly through the pits, where it does not to cross thick secondary walls. Vessel elements are generally wider, shorter, thinner walled, and less tapered than the tracheids. b. The end walls between sieve-tube elements, called sieve plates, haw pores that facilitate the flow of fluid from cell to cell along the sieve tube. Alongside each sieve- tube element is a nonconducting cell called a companion which is connected to the sieve-tube element by numerous channels, plasmodesmata 16. summary charts. a. The three plant organs are : roots, stems, and leaves b. The three basic plant tissues are: dermal, vascular, ground c. The three basic plant cells are: Parenchyma, Collenchyma, Sclerynchyma, xylem and phloem 17. What is the difference between indeterminate growth and determinate growth?

Sieve plate with pores (SEM)

A major difference between plants and most animals is that plant growth is not limited to an embryonic or juvenile period. Instead, growth occurs throughout the plant's life, a process known as indeterminate growth. At any given time, a typical plant consists of embryonic, developing, and mature organs. Except for dormant periods, most plants grow continuously. In contrast, most animals and some plant organs-such as most leaves, thorns, and flowers-undergo determinate growth; that is, they stop growing after reaching a certain size. 18. Although plants generally show indeterminate growth, what are three examples of plant parts that show determinate growth? Leaves, thorns, and flowers 19. Based on the length of their life cycle, plants are categorized into three groups. Explain what each category means below, and provide an example. a. Annuals complete their life cycle-from germination to flower ing to seed production to death-in a single year or less. Many wildflowers are annuals, as are most staple food crops, including legumes and cereal grains such as wheat and rice. b. Biennials generally require two growing seasons to complete their life cycle, flowering and fruiting only in their second year. Radishes and carrots are biennials, though they are usually harvested after the first year.

al bud scale

ry buds

r;;::::;
One-year-old side branch formed from aXillary bud near shoot tip

scar

scar left by apical scales of previous ters

Perennials live many years and include trees, shrubs, and some grasses. Some buffalo grass of the North American plains is thought to have been growing for 10,000 years from seeds that sprouted at the close of the last ice age. 20. Plants are capable of indeterminate growth because they have perpetually embryonic tissues called meristems. 21. Explain the following relationships: lengthens a. Apical meristems and primary growth: Apical meristems, located at the tips of roots and shoots and in the axillary buds offshoots, provide additional cells that enable roots and shootso grow in length, a process known as primary growth. Primary growth the plant t As you have learned, primary growth is xtend inhroughout the soil and shoots to increase their exposure to allows roots to egrowth t length, produced by apical meristems.hCollectively, thenonwoody) plants, primary growth produces all, or almost all light. In erbaceous ( results of this growth are calledof the plant plant body. In herbaceous the primary body. plants, it is usuallyLateral meristems and secondary growth: Woody plants. however, also grow in b. the entire plant. In woody plants, it consists only of the youngest parts,n the parts of stems Although girth i which are not yet woody. and roots that no longer grow in length. This growth in apical meristems lengthen both roots and shoots, there are thickness, known as secondary growth, is caused by the activity of lateral meristems differences in the primary growth of these two systems. called the vascular cambium and cork cambium. c. Primary growth and secondary growth: Primary growth allows roots to extend Primary Growth of Roots throughout the soil and shoots to increase their exposure to light. In herbaceous The tip of a root is covered by a thimble-likeprimary which (nonwoody) plants, root cap, growth produces all, or almost all. of the plant body. protects the delicate apical lants. however, also grow in girth in the parts of stems and roots that no Woody p meristem as the root pushes through the abrasive soil during primary growth. The root cap in thickness, known as secondary growth. longer grow in length. This growth also secretes a polysaccharide slime that lubricates the soil 22. The figure below shows an image that is like a slide many students study in a mitosis lab and around the tip ohhe root. Growth occurs just behind the tip is labeled for this lesson as the Primary growth of a root. Label the nine structures shown in three zones of cells at successive stages of primary growth. in the figure: cortex, vascular cylinder, epidermis, apical meristem, root cap, root hair, zone Moving away from the tip, they are the zones of cell division, of d and differentiation one o 35.13). elongation, ifferentiation, z(Figuref elongation, and zone of cell division.
Cortex K.y to labels Dermal Ground Vascular Vascular cylinder

c.

h in a winter twig.

sed by scales that protect n spring, the bud sheds its mary growth, producing a ng each growth segment, ere left when leaves fell. ud or a branch formed by wig are bud scars from the ical bud during the previason, primary growth exrowth thickens the parts

Zone of differentiation

Zone of elongation

nd secondary growth. n divide and replace rface. Why is it inacof cell division to a 100llm

Zone of cell division

inately, but leaves do plant? er picks some radishes l. Since radishes are e remaining plants in will grow larger during idea? Explain.


... Figure 35.13 Primary growth of a root. The light micrograph and diagram reveal the anatomical features of the tip n the zone of cell division, zone of elongation, and zone of 23. Explain what events occur i of an ooion root. Mitosis is concentrated in the zone of cell dil'ision, where the apical differentiation. menstem and its immediate growth products are located. The apical menstem also maintains the root cap by generating new cells that replace those that occurs The three are sloughedrade together, of the root o zonein theboundaries. The zone of cell division includes the root zones g off. Most lengthening with the zone of elongation. Cells become functionally mature inn sharp of apical meristem and its derivatives. New root cells are produced in this region, including the root cap. differentiatioo. The zones grade into one another without sharp boundaries.

x A.

Typically, about a millimeter behind the tip of the root is the zone of elongation, where root cells
CIlAPTE TIlIRlYfIVE

Plant Structure, Growth, and Development

747

----cylinder ----Pericycle Core of parenchyma cells

elongate, sometimes to more than ten times their original length. Cell elongation in this zone pushes the tip farther into the soil. Meanwhile, the root apical mXylem eristem keeps adding cells to the younger f-; end of the zone of elongation. Even before the root cells finish lengthening, many begin specializing 100llm Phloem in structure and function. In the zone of differentiation, or zone of maturation, cells complete their differentiation and become phloem in cell center (a) Root with xylem and distinct the types.

f-;

100llm

(typical of eudicots). In the roots of typical gymnosperms and (b) issue located in the 24. In most roots, teudicots. as well as hloem is a solid cylinder of vascular tRoot with parenchyma in the center (typical of he xylem and psome monocots, the stele is a vascular cylinder monocots). The stele of many monocot roots center of the root and called the stele. The figure below shows the sis a vasculardicot root. Label parenchyma tele of a cylinder with a core of consisting of a lobed core of KYIem with the xylem, phloem, endodermis, and pericycle. Also define the two new terms as indicated. a ring of phloem phloem between the lobes. surrounded by a ring of KYIem and

to labels Pericycle Dermal Ground Vascular

:L'lt7Tt---1Phloem

... Figure 35.14 Organization of primary tissues in young roots. Parts (a) and (b) show cross sections of the roots of Ranuncu/us (buttercup) and lea (maize), respectively. These Endodermis: The innermost layer of the cortex is called the endodermis, a cylinder one cell thick that represent two basic patterns of root organization. of which there are many variations, depending forms the boundary with the vIlm 50 ascular cylinder. on the plant species (alllMs).

Pericycle: Lateral roots arise from the pcricyclc, the outermost cell layer in the vascular cylinder, 748 UNIT SIX Plant Form and nside t which is adjacent to and just iFunctionhe endodermis. 25. Why must new roots formed by the pericycle originate in the center of the root? A lateral root cannot originate near the root's surface because its vascular system must be continuous with the vascular cylinder at the center of the established root. 26. From Figure 35.16, label shoot apical meristem, leaf primordia, young leaf, developing vascular strand, and axillary bud meristems.

lateral root. A lateral root originates in the pericyde, of a root, and grows out through the and s, the view of the original root is a cross section. while section. Shoot apical meristem leaf primordia

ng mostly ofparenchyma een the vascular cylinder nd tissue store carbohybsorb water and minerals f the cortex is called the that forms the boundary see in Chapter 36, the enegulates passage of subcylinder. yclc, the outermost cell adjacent to and just inside teral root pushes through rges from the established annot originate near the tem must be continuous of the established root.

Young leaf

Developing vascular strand

bud meristems

shaped mass of dividing Leaves develop from leaf 0.25 mm finger-like projections .. Figure 35.16 The shoot tip. Leal primordia arise from the m. Axillary buds develop 27.flanks of the dome of ithethis figure is responsible for primary growth? What structure n apical meristem. This is a longitudinal section of the shoot tip of Coleus (LM). t by the apical meristem Apical meristem illary buds can form latTissue Organization of Stems gure 35.12). 28.Theis possible to tell a young eudicot from a monocot by the structure of the stem. In the It epidermis covers stems as part of the continuous dermal paced close together befollowing figure, label the eudicot, monocot, epidermis, and vascular bundles. tissue system. Vascular tissue runs the length of a stem in vasMost shoot elongation is cular bundles. Unlike lateral roots, which arise from vascular lls below the shoot tip. In tissue deep within a root and disrupt the vascular cambium, eafcells are produced by Phloem Xylem cortex, and epidermis as they emerge (see Figure 35.15), latd from the apical merieral shoots develop from axillary bud meristems on the stem's meristems, remain at the surface and disrupt no other tissues (see Figure 35.16). The des. This morphological 9 because the elevated part vascular bundles ofthe stem converge with the root's vascular 9 out stopping growth. cylinder in a zone of transition located near the soil surface. Q) I> '0 <l>
CIlAPTE TIlIRlYfIVE

Plant Structure, Growth, and Development


Pith Epidermis
K.y to labels

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..

., " ..

'"

co

<l>

e.
Vascular
bundles

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'"
f------<

Epidermis

Vascular bundle
1 mm
(3) Cross section of stem with vascular bundles forming a ring (typical of eudicots). Ground tissue toward the

Dermal

Ground

Vascular

1 mm (b) Cross section of stem with scattered vascular bundles

(typical of monaco!s). In such an arrangement, ground tissue is

inside is called pith, and ground tissue toward the outside is

not partitioned into pith and cortex (LM).

called cortex (lM).

29. How is the arrangement of vascular bundles different in monocot and dicot stems? In mcular bundles arranged in a ring t(figureascular tissue consists of vascular bof many eudicots have twoin a ring. The xylem ost eudicot species, he v 35.17a). The xylem photosynthesis. The leaves undles arranged disin each vascular bundle is adjacent to the pith, and the phloem in each bundle is adjacent to the cortex. In most monocot stems, the vascular bundles are scattered throughout the ground tissue, rather than forming a ring (Figure 35.17b). In the stems of both monocots and eudicots, the ground tissue consists mostly of parenchyma cells. However, collenchyma cells just beneath the epidermis strengthen many stems. Scle renchyma cells, especially fiber cells, also provide support in tinct areas: palisade mesophyll and spongy mesophyll. The In most eudicot species, the vascular tissue consists of vasophyll consists mainly of parenchyma cells specialized for

II

.... Figure 35.17 Organization of primary tissues in young stems. Why aren't the terms pith and cortex used to describe the ground tissue of monocot stems?

palisade mesophyll consists of one or more layers of elongated parenchyma cells on the upper part of the leaf. The

spongy mesophyll is below the palisade mesophyll. These


parenchyma cells are more loosely arranged, with a labyrinth of air spaces through which CO 2 and oxygen circulate around the cells and up to the palisade region. The air spaces are particularly large in the vicinity of stomata, where gas ex-

in each vascular bundle is adjacent to the pith, and the phloem in each bundle is adjacent to the cortex. In most monocot stems, the vascular bundles are scattered throughout the ground tissue, rather than forming a ring. 30. To understand the process of photosynthesis, students are expected to know leaf structure in greater detail. Using Figure 35.18, label each structure just as shown in the text.
Figure 35.18 Leafanatomy.
Guard cells

K.y
to labels

Dermal Ground Vascular _ .... Sderenchyma .... fibers

(b) Surface view of a spiderwort (rradescantia) leaf (LM)

Upper epidermis Palisade mesophyll

Bundlesheath cell

t
CHECK

Spoogy mesophyll
Lower epidermis

(a) Cutaway drawing of leaf tissues

the previous layer (see Figure 35.19). In this way, the vascular leaves. In contrast. secondary growth, the growth in thickness cambium thickens roots and stems. The stem thickens as the vascular cambium forms secondary to woodynside and secondary pconsists ofo produced by lateral meristems, occurs in stems and roots of In a typical the i stem, the vascular cambium hloem t the outside. but rarely in leaves. The secondary plant body woody plants, a continuous cylinder ofundifferentiated parenchyma cells, loconsists of the tissues produced by the vascular cambium and cated outside the pith and primary xylem and to the inside cork cambium.hat cells are formed to the inside and the outside of the vascular cIn a typical woody root, the of the cortex and primary phJoem. ambium? b. W The vascular cambium adds secondary xylem (wood) and secondary phloem, increasing vascular flow and supvascular cambium forms to the exterior of the primary xylem port for the shoot system. The cork cambium produces a tough, and interior to the primary phloem and pericyde.
CIlAPTE TIlIRTYfIVE

thick covering consisting mainly of wax-impregnated ceUs that 35.3 protect through s ..','ater loss and 31. What g roots and shoots differ in branching. I. Describe howas critical to photosynthesis enters the leaf the stem fromtoma? CO2 from invasion by insects, bacteria, and fungi. All ater 32. What is lost through t and toma 2. Contrast primary growth in rootshe sshoots. that leads to transpiration? Wgymnosperm species and many eudicot species have secondary growth, but it is rare in monocots. 33. I grazing animals are removed leaf? C4 3. Whens this a C3, C4, or CAM from grasslands, Primary growth and secondary growth occur eudicots often replace grasses. Suggest a reason why. 34. Primary growth arises from apical meristems and results in ___growth___ of roots, simultane- nd stems, a ously. As primary growth adds leaves and lengthens stems and 4, _i,ij:f.jlijM If econdary growth arises from _____________ and ___vascular_____ cambium and results a leaf is vertically oriented, would you leaves. S expect its mesophyll to__girth___ of spongy and stems. roots in the younger regions of a plant, secondary growth in increased _ be divided into roots and thickens stems and roots in older regions where primary palisade layers? Explain. 35. Explain what is produced by these structures growth has stopped. The process is similar in shoots and For suggested answers. see Appendix A. a. Vascular cambium: The vascular cambium is a35.19, on theonext page, provides an overview ofen roots. Figure cylinder f meristematic cells, of- t only one cell thick. It increases in circumference and also adds layers of secondary growth in a woody stem. xylem to its interior and secondary phloem to its exterior. The Vascular Cambium and Secondary b. Cork Cambium: During the early stages of secondary growth, the epidermis is Vascular Tissue pushed outward, causing it to split, dry, and fall off the stem or root. It is replaced by adds girth tto first cork cvascular cambium is a cylinder of meristematic cells, ofThe ambium, a cylinder of dividing cells that two tissues produced by he ten n t one uter layer of the ericycle in and also stems andarises in in woody plantstems and ionlyhe ocell thick. It increases inpcircumference roots. roots the outer cortex of s adds layers of secondary xylem to its interior and secondary 36. Read the text that accompanies Figure 35.19 and then answer these questions. As you have seen, primary growth arises from apical meristems phloem to its exterior. Each layer has a larger diameter than a. What results in primary growth of the stem? and involves the production and elongation of roots, stems, and
CONCEPT

Plant Structure, Growth, and Development

751

---+ y--+-

Primary Vascular cambium Primary phloem

Epidermis

as the vascular cambium forms secondary to the inside and secondary phloem to the outside.

y--+-

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As the Vascular cambium's diameter increases. The secondary phloem and other tissues to vascular "y As the vascular cambium's diameter the cambium can't keep pace because their cells no longer divide, As a result, these tissues, increases. the secondary phloem and including the epidermis, will eventually rupture. A second lateral mcambium the cork eristem, other tissues to the cambium, develops from parenchyma cells in the keep paceThe cork cambium produces cork can't cortex. because their cells no longer divide, As a result, these cells, which replace the epidermis. Primary tissues, including the epidermis, will rylem c. What is the difference in the formation of peventuallyxrupture.aA second lateral ersus secondary rimary ylem nd phloem v meristem, the cork cambium, develops Secondary rylem xylem and phloem? from parenchyma cells in the cortex. Vascular cambium Primary xylem is the xylem that is formed during primary growth from cells, The cork cambium produces cork procambium. It Secondary phloem which evelops fter the includes protoxylem and metaxylem. Metaxylem dreplace theaepidermis. protoxylem but before secondary xylem. It is distinguished by wider vessels and tracheids. As it develops, the xylem { Pflmary phloem First cork cambium Cork can become endarch or exarch. 01n year 2 of secondary growth, the vascular secondary growth Secondary xylem is the xylem that is formed during cambium produces morefrom vascular cambium. Although secondary xylem is also fsecondary members of the "gymnosperm" groups ound in xylem and phloem, and the cork cambium produces more cork. Gnetophyta and Ginkgophyta and to a lesser extent in members of the Cycadophyta As the stem's diameter increases. the outermost unction of to the cork 37. What vascular tissue forms the bark, and what is the ftissues exteriorthe bark?

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Some initials of the vascular cambium give rise to vascular rays (see page),

cambium rupture and are sloughed off,

Many people think that bark is only the protective outer covering of a woody stem or root. Actually, In many cases, the cork cambium bark includes all tissues external to the vascular cambium. In an outward direction, its main components are the secondary phloem (produced by re-forms deeper cambium), When ost recent the vascular in the corteK. the m Secondary none of the cortex is left, the cambium Kylem (two periderm, and all the older layers of periderm develops from phloem parenchyma cells.

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38. What tissues are included in the bark of a tree? Vascular cambium Secondary Phloem and layers of periderm Each cork cambium and the tissues it Secondary phloem produces form a layer of periderm, OMost recent 39. On this figure, add these labels: cork cambium, cork, periderm, bark, growth ring, secondary Bark consists of all tissues exterior to cork cambium Cork xylem, secondary phloem, and vascular cambium. the vascular cambium.

years of production)

Secondary phloem Vascular cambium Secondary wood Early wood

em' } Periderm cambium


Cork

35.19 Primary and secondary of a stem. The progress of secondary n be tracked by the sections equentially older parts of the stem, does the vascular cambium cause some s to rupture?

O.5mm

Vascular ray Growth ring (b) Cross section of a three-yearold Tilia (linden) stem (LM)

40. Look back at the stem in Figure 35.19 and find the horizontal slits in the bark, known as lenticels. You may have noticed lenticels on the young twigs of trees or shrubs. What is the function of lenticels?
Plant Form and Function

Dotting the peridem are small, raised areas called lenticels, in which there is more space between cork cells, enabling living cells within a woody stem or root to exchange gases with the outside air. Lenticels often appear as slits.