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Chapter 5
Cassava Botany and Physiology

Alfredo Augusto Cunha Alves

Embrapa Cassava and Fruits, Caixa Postal 007, 44.380–000, Cruz das Almas,
Bahia, Brazil

Introduction cuttings. The seedlings are genetically segre-

gated into different types due to their
Cassava is a perennial shrub of the family reproduction by cross-pollination. If propagated
Euphorbiaceae, cultivated mainly for its starchy by cuttings under favourable conditions, sprout-
roots. It is one of the most important food staples ing and adventitious rooting occur after 1 week.
in the tropics, where it is the fourth most impor-
tant source of energy. On a worldwide basis it
is ranked as the sixth most important source of Morphological and Agronomic
calories in the human diet (FAO, 1999). Given Characteristics
the crop’s tolerance to poor soil and harsh clima-
tic conditions, it is generally cultivated by small Cassava, which is a shrub reaching 1–4 m
farmers as a subsistence crop in a diverse range height, is commonly known as tapioca, manioc,
of agricultural and food systems. Although mandioca and yuca in different parts of the
cassava is a perennial crop, the storage roots can world. Belonging to the dicotyledon family
be harvested from 6 to 24 months after planting Euphorbiaceae, the Manihot genus is reported to
(MAP), depending on cultivar and the growing have about 100 species, among which the only
conditions (El-Sharkawy, 1993). In the humid commercially cultivated one is Manihot esculenta
lowland tropics the roots can be harvested after Crantz. There are two distinct plant types: erect,
6–7 months. In regions with prolonged periods with or without branching at the top, or
of drought or cold, the farmers usually harvest spreading types.
after 18–24 months (Cock, 1984). Moreover, The morphological characteristics of
the roots can be left in the ground without cassava are highly variable, which indicate a
harvesting for a long period of time, making it high degree of interspecific hybridization. There
a very useful crop as a security against famine are many cassava cultivars in several germplasm
(Cardoso and Souza, 1999). banks held at both international and national
Cassava can be propagated from either stem research institutions. The largest germplasm
cuttings or sexual seed, but the former is the bank is located at Centro Internacional de
commonest practice. Propagation from true seed Agricultura Tropical (CIAT), Colombia, with
occurs under natural conditions and is widely approximately 4700 accessions (Bonierbale
used in breeding programmes. Plants from true et al., 1997), followed by EMBRAPA’s collection
seed take longer to become established, and they in Cruz das Almas, Bahia, with around 1700
are smaller and less vigorous than plants from accessions (Fukuda et al., 1997), representing

©CAB International 2002. Cassava: Biology, Production and Utilization

(eds R.J. Hillocks, J.M. Thresh and A.C. Bellotti) 67

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68 A.A.C. Alves

the germplasm of the following Brazilian eco- buds under the soil. These roots develop to make
systems: lowland and highland semiarid tropics, a fibrous root system. Only a few fibrous roots
lowland humid subtropics, lowland subhumid (between three and ten) start to bulk and become
tropics, lowland humid tropics, and lowland hot storage roots. Most of the other fibrous roots
savannah. The cassava genotypes are usually remain thin and continue to function in water
characterized on the basis of morphological and and nutrient absorption. Once a fibrous root
agronomic descriptors. Recently, the Interna- becomes a storage root, its ability to absorb water
tional Plant Genetic Resources Institute (IPGRI) and nutrients decrease considerably. The storage
descriptors (Gulick et al., 1983) were revised, and roots result from secondary growth of the fibrous
a new version was elaborated, in which 75 roots; thus the soil is penetrated by thin roots,
descriptors were defined, 54 being morpho- and their enlargement begins only after that
logical and 21 agronomic (Fukuda et al., 1997). penetration has occurred.
Morphological descriptors (for example, lobe Anatomically, the cassava root is not a
shape, root pulp colour, stem external colour) tuberous root, but a true root, which cannot
have a higher heritability than agronomic (such be used for vegetative propagation. The mature
as root length, number of roots per plant and root cassava storage root has three distinct tissues:
yield). Among morphological descriptors, the bark (periderm), peel (or cortex) and paren-
following were defined as the minimum or basic chyma. The parenchyma, which is the edible
descriptors that should be considered for identify- portion of the fresh root, comprises approxi-
ing a cultivar: (i) apical leaf colour; (ii) apical leaf mately 85% of total weight, consisting of xylem
pubescence; (iii) central lobe shape; (iv) petiole vessels radially distributed in a matrix of starch-
colour; (v) stem cortex colour; (vi) stem external containing cells (Wheatley and Chuzel, 1993).
colour; (vii) phyllotaxis length; (viii) root ped- The peel layer, which is comprised of
uncule presence; (ix) root external colour; (x) sclerenchyma, cortical parenchyma and phloem,
root cortex colour; (xi) root pulp colour; (xii) root constitutes 11–20% of root weight (Barrios
epidermis texture; and (xiii) flowering. and Bressani, 1967). The periderm (3% of total
Given the large number of cassava genotypes weight) is a thin layer made of a few cells thick
cultivated commercially and the large diversity and, as growth progresses, the outermost por-
of ecosystems in which cassava is grown, it is tions usually slough off. Root size and shape
difficult to make a precise description of the mor- depend on cultivar and environmental condi-
phological descriptors as there is a genotype- tions; variability in size within a cultivar is
by-environmental conditions interaction. Thus, greater than that found in other root crops
in addition to morphological characterization, (Wheatley and Chuzel, 1993). Table 5.1 lists
molecular characterization, based mainly on morphological and agronomic characteristics of
DNA molecular markers, has been very useful in the root and its variability in cassava.
order to evaluate the germplasm genetic diver-
sity (Beeching et al., 1993; Fregene et al., 1994).
Stems

Roots The mature stem is woody, cylindrical and

formed by alternating nodes and internodes. On
Roots are the main storage organ in cassava. the nodes of the oldest parts of the stem, there
In plants propagated from true seeds a typical are protuberances, which are the scars left by
primary tap root system is developed, similar the plant’s first leaves. A plant grown from stem
to dicot species. The radicle of the germinating cuttings can produce as many primary stems as
seed grows vertically downward and develops there are viable buds on the cutting. In some
into a taproot, from which adventitious roots cultivars with strong apical dominance, only
originate. Later, the taproot and some adventi- one stem develops.
tious roots become storage roots. The cassava plant has sympodial branch-
In plants grown from stem cuttings the roots ing. The main stem(s) divide di-, tri- or tetra-
are adventitious and they arise from the basal- chotomously, producing secondary branches
cut surface of the stake and occasionally from the that produce other successive branchings. These

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Botany and Physiology 69

Table 5.1. Some morphological and agronomic characteristics of roots and their variability in cassava.

Root characteristic Variability Reference

Morphological
External colour White or cream; yellow; light Fukuda and Guevara (1998)
brown; dark brown
Cortex colour White or cream; yellow; pink; purple Fukuda and Guevara (1998)
Pulp (parenchyma) colour White; cream; yellow; pink Fukuda and Guevara (1998)
Epidermis texture Smooth; rugose Fukuda and Guevara (1998)
Peduncule Sessile; pedunculate; both Fukuda and Guevara (1998)
Constriction None or little; medium; many Fukuda and Guevara (1998)
Shape Conical; conical–cylindrical; Fukuda and Guevara (1998)
cylindrical; irregular
Agronomic
No. storage roots/plant 3–14 Dimyati (1994); Wheatley and
Chuzel (1993); Ramanujam and
Indira (1983); Pinho et al. (1995)
Weight of storage roots/plant 0.5–3.4 kg FW Dimyati (1994); Wheatley and
Chuzel (1993); Ramanujam and
Indira (1983)
Weight of one storage root 0.17–2.35 kg Barrios and Bressani (1967);
Ramanujam and Indira (1983)
Length of storage root 15–100 cm Wheatley and Chuzel (1993);
Barrios and Bressani (1967);
Pinho et al. (1995)
Diameter of storage root 3–15 cm Wheatley and Chuzel (1993);
Barrios and Bressani (1967);
Pinho et al. (1995)
Diameter of fibrous root 0.36–0.67 mm Connor et al. (1981)
Depth of fibrous root Up to 260 cm Connor et al. (1981)
Amylose in root starch 13–21% FW O’Hair (1990)
Protein in whole root 1.76–2.68% FW Barrios and Bressani (1967)
Protein in pulp (parenchyma) 1.51–2.67% FW Barrios and Bressani (1967)
1.0–6.0% DW Wheatley and Chuzel (1993)
Protein in peel 2.79–6.61% FW Barrios and Bressani (1967)
7.0–14.0% DW Wheatley and Chuzel (1993)
DM in whole fresh root 23–43% Barrios and Bressani (1967);
Ghosh et al. (1988); Ramanujam
and Indira (1983); O’Hair (1989)
DM in peel 15–34% Wheatley and Chuzel (1993);
Barrios and Bressani (1967);
O’Hair (1989)
DM in pulp 23–44% Wheatley and Chuzel (1993);
Barrios and Bressani (1967);
O’Hair (1989)
Carbohydrates in whole root 85–91% DW Barrios and Bressani (1967)
Carbohydrates in peel 60–83% DW Barrios and Bressani (1967)
Carbohydrates in pulp 88–93% DW Barrios and Bressani (1967)
(parenchyma)
Starch in whole root 20–36% FW Wholey and Booth (1979);
O’Hair (1989); Ternes et al.
(1978)
77% DW Ghosh et al. (1988)
Starch in peel 14–25% FW O’Hair (1989)
44–59% DW Wheatley and Chuzel (1993)
Continued

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70 A.A.C. Alves

Table 5.1. Continued.

Root characteristic Variability Reference

Starch in pulp (parenchyma) 26–40% FW O’Hair (1989)

70–91% DW Wheatley and Chuzel (1993)
Peel in whole root 11–20% FW Barrios and Bressani (1967)
Crude fibre in whole root 3.8–7.3% DW Barrios and Bressani (1967)
Crude fibre in peel 9.2–21.2% DW Barrios and Bressani (1967)
5.0–15.0% DW Wheatley and Chuzel (1993)
Crude fibre in pulp 2.9–5.2% DW Barrios and Bressani (1967)
(parenchyma) 3.0–5.0% DW Wheatley and Chuzel (1993)
Total sugars 1.3–5.3% DW Wheatley and Chuzel (1993)

DM, dry matter; FW, fresh weight; DW, dry weight.

branchings, which are induced by flowering, 0.5–1.0 cm long), which remain attached to the
have been called ‘reproductive branchings’. stem when the leaf is completely developed
Stem morphological and agronomic char- (CIAT, 1984). The petiole length of a fully opened
acteristics are very important to characterizing leaf normally varies from 5 to 30 cm, but may
a cultivar (Table 5.2). The variation of these reach up to 40 cm.
characteristics depends on cultivar, cultural The upper leaf surface is covered with a
practice and climatic conditions. shiny, waxy epidermis. Most stomata are located
on the lower (abaxial) surface of the leaves;
only a few can be found along the main vein on
Leaves the upper (adaxial) surface (Cerqueira, 1989). Of
1500 cultivars studied, only 2% had stomata
Cassava leaves are simple, formed by the lamina on the adaxial surface (El-Sharkawy and Cock,
and petiole. The leaf is lobed with palmated 1987a). The stomata on the upper surface are
veins. There is generally an uneven number of also functional and bigger than those on the
lobes, ranging from three to nine (occasionally undersurface. Both are morphologically para-
11). Only a few cultivars are characterized by cytic, with two small guard cells surrounded
having three-lobed mature vegetative leaves, by two subsidiary cells (Cerqueira, 1989). The
which may represent the primitive ancestral number of stomata per leaf area range from
form (Rogers and Fleming, 1973). Leaves near 278 to 700 mm−2, and all stomatal pores can
the inflorescence are generally reduced in size occupy from 1.4 to 3.1% of the total leaf area
and lobe number (most frequently three-lobed), (Table 5.3).
but the one closest to the base of the inflores-
cence is frequently simple and unlobed.
Leaves are alternate and have a phyllotaxy Flowers
of 2/5, indicating that from any leaf (leaf 1) there
are two revolutions around the stem to reach the Cassava is a monoecious species producing both
sixth (leaf 6) in the same orthostichy as leaf 1. In male (pistillate) and female (staminate) flowers
these two revolutions there are five successive on the same plant. The inflorescence is generally
intermediate leaves (not counting leaf 1). formed at the insertion point of the reproductive
The main leaf morphological and agro- branchings; occasionally inflorescences can be
nomic characteristics and their variation are found in the leaf axils on the upper part of the
given in Table 5.3. Many of them (mainly the plant. The female flowers, located on the lower
morphological ones) are used to characterize part of the inflorescence, are fewer in number
cultivars and may vary with environmental than male flowers, which are numerous on the
conditions and plant age. upper part of the inflorescence. On the same
Mature leaves are glabrous and each leaf inflorescence, the female flowers open 1–2
is surrounded by two stipules (approximately weeks before the male flowers (protogyny).

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Table 5.2. Some morphological and agronomic characteristics of stems and their variability in cassava.

Stem characteristic Variability Reference

Morphological
Cortex (collenchyma) colour Yellow; light green; dark green Fukuda and Guevara (1998)
External colour Orange; green–yellow; gold; dark Fukuda and Guevara (1998)
brown; silver; grey; dark brown
Phyllotaxis length Short (< 8 cm); medium (8–15 cm); Fukuda and Guevara (1998)
large (> 15 cm)
Epidermis colour Cream; light brown; dark brown; Fukuda and Guevara (1998)
orange
Growth habit Straight; zigzag Fukuda and Guevara (1998)
Apical stem colour Green; green-purple; purple Fukuda and Guevara (1998)
Branching habit Erect; dichotomous; trichotomous; Fukuda and Guevara (1998)
tetrachotomous
Agronomic
Diameter of mature stem 2–8 cm CIAT (1984); Ramanujam and
Indira (1983)
Plant height 1.20–3.70 m Ramanujam and Indira (1983);
Ramanujam (1985); Veltkamp
(1985a); Pinho et al. (1995)
No. of nodes from planting– 22–96 Veltkamp (1985a)
1st branch level
No. of days from planting– 49–134 Veltkamp (1985a)
1st branch level
No. of apices/plant 2.8–27.5 Pinho et al. (1995)

Male and female flowers on different branches three undulated, fleshy lobes originates from the
of the same plant can open at the same time. style.
Normally, cassava is cross-pollinated by insects;
thus it is a highly heterozygous plant.
The flowers do not have a calyx or corolla, Fruit and seeds
but an indefinite structure called perianth or
perigonium, made up of five yellow, reddish or The fruit is a trilocular capsule, ovoid or globu-
purple tepals. The male flower is half the size of lar, 1–1.5 cm in diameter and with six straight,
the female flower. The pedicel of the male flower prominent longitudinal ridges or aristae. Each
is thin, straight and very short, while that of the locule contains a single carunculate seed. The
female flower is thick, curved and long. Inside the fruit has a bicidal dehiscence, which is a combi-
male flower, there is a basal disk divided into ten nation of septicidal and loculicidal dehiscences,
lobes. Ten stamens originate from between them. with openings along the parallel plane of the
They are arranged in two circles and support the dissepiments and along the midveins of the
anthers. The five external stamens are separated carpels, respectively. With this combination of
and longer than the inner ones, which join dehiscences, the fruits open into six valves caus-
together on the top to form a set of anthers. The ing an explosive dehiscence, ejecting the seeds
pollen is generally yellow or orange, varying some distance (Rogers, 1965). Fruit maturation
from 122 to 148 µm in size, which is very large generally occurs 75–90 days after pollination
compared to other flowering plants (Ghosh et al., (Ghosh et al., 1988). The seed is ovoid–
1988). The female flower also has a ten-lobed ellipsoidal, approximately 100 mm long, 6 mm
basal disk, which is less lobulated than the male wide and 4 mm thick. The weight varies from 95
flower. The ovary is tricarpellary with six ridges to 136 mg per seed (Ghosh et al., 1988). The
and is mounted on the basal disk. The three smooth seed coat is dark brown, mottled with
locules contain one ovule each. A very small style grey. The seeds usually germinate soon after col-
is located on top of the ovary, and a stigma with lection, taking about 16 days for germination.

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72 A.A.C. Alves

Table 5.3. Some morphological and agronomic characteristics of leaves and their variability in cassava.

Leaf characteristic Variability Reference

Morphological
Apical leaf colour Light green; dark green; green–purple; Fukuda and Guevara (1998)
purple
Apical pubescence Absent; present Fukuda and Guevara (1998)
Shape of central lobe Ovoid; elliptic–lanceolate; obovate– Fukuda and Guevara (1998)
lanceolate; oblanceolate; lanceolate;
linear; pandurate; linear–pyramidal;
linear–pandurate; linear–hostatilobada
Petiole colour Green–yellow; green; green–red; Fukuda and Guevara (1998)
red-green; red; purple
Mature leaf colour Light green; dark green; green-purple; Fukuda and Guevara (1998)
purple
Protuberance of leaf scars No protuberance; protuberant Fukuda and Guevara (1998)
No. of lobes 3; 5; 7; 9; 11 Fukuda and Guevara (1998)
Agronomic
Petiole length 5–30 cm Ghosh et al. (1988)
9–20 cm CIAT (1984)
Total chlorophyll 2.18–2.86 mg g−1 leaf FW Ramanujam and Jos (1984)
Central lobe length 4–20 cm CIAT (1984)
Central lobe width 1–6 cm CIAT (1984)
No. of stomata/leaf area 278–700 mm−2 Ghosh et al. (1988); Cerqueira
in adaxial epidermis (1989); Splittstoesser and Tunya
(1992); Connor and Palta (1981)
Relative area of stomata 1.4–3.1% Cerqueira (1989); Pereira and
pore (% from leaf area) Splittstoesser (1990)
Stipule length 0.5–1.0 cm CIAT (1984)
Leaf thickness 100–120 µm Pereira and Splittstoesser (1990)
DM in mature leaf 25% Barrios and Bressani (1967)
Fibre in mature leaf 4.58% DW Barrios and Bressani (1967)
Ash in mature leaf 8.28% DW Barrios and Bressani (1967)
Protein in mature leaf 7.1–8.9% FW Barrios and Bressani (1967)
28.8% DW Barrios and Bressani (1967)
Soluble carbohydrates 11.36% FW Barrios and Bressani (1967)
44.84% DW Barrios and Bressani (1967)

DM, dry matter; FW, fresh weight; DW, dry weight.

Growth and Development depend on several factors related to varietal dif-

ferences, environmental conditions and cultural
Plant developmental stages practices. The initial growth (at 15-day inter-
vals) from emergence to 150 days is presented in
As cassava is a perennial shrub it can grow Fig. 5.2. Growth at 60-day intervals during the
indefinitely, alternating periods of vegetative first cycle (0–360 days after planting; DAP) is
growth, storage of carbohydrates in the roots, shown in Fig. 5.3. The results in these two fig-
and even periods of almost dormancy, brought ures are consistent with other authors (Howeler
on by severe climatic conditions such as low and Cadavid, 1983; Ramanujam and Biradar,
temperature and prolonged water deficit. There 1987; Távora et al., 1995; Peressin et al., 1998).
is a positive correlation between the total The periods and main physiological events
biomass and storage root biomass (Fig. 5.1; during the growth of a cassava plant under
Ramanujam, 1990). During its growth, there favourable conditions in the field can be
are distinct developmental phases. The occur- visualized in these figures and are summarized
rence, duration and existence of each phase below:

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Botany and Physiology 73

Fig. 5.1. Relationship between dry weight (DW) of whole plant (x) and DW of storage roots (y) for
individual plants of a field trial at the University of the West Indies: y = 0.56x−34; r 2 = 0.96; n = 112.
(Source: Boerboom, 1978.)

Fig. 5.2. Partitioning of dry matter during the initial development of cassava cv. Cigana, Cruz das
Almas, Bahia, Brazil. (Source: Porto, 1986.)

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74 A.A.C. Alves

Fig. 5.3. Growth of cassava plant during the first cycle (12 months). Average of two varieties. DAP,
days after planting. Graph made from data of Lorenzi (1978).

Emergence of sprouting – 5–15 DAP storage roots represent 10–15% of total dry
matter (DM; Fig. 5.2).
• From 5–7 DAP the first adventitious roots
arise from the basal cut surface of the stake
and occasionally from the buds under the Development of stems and leaves (canopy
soil. establishment) – 90–180 DAP
• 10–12 DAP the first sprouting occurs,
• Maximum growth rates of leaves and stems
followed by small leaves which start to
are achieved in this period, and the branch-
emerge (Conceição, 1979).
ing habit and plant architecture is defined
• Emergence is achieved at 15 DAP.
(Fig. 5.3).
Beginning of leaf development and formation
• From 120 to 150 DAP the leaves are able to
intercept the most of the incident light on
of root system – 15–90 DAP
canopy (Veltkamp, 1985c).
• The true leaves start to expand around 30 • Maximum canopy size and maximum DM
DAP (Fig. 5.2) when the photosynthetic partition to leaves and stems are accom-
process starts to contribute positively to plished (Howeler and Cadavid, 1983;
plant growth. Ramanujam, 1985; Távora et al., 1995).
• Until 30 DAP, shoot and root growth • The storage root continues to bulk.
depends on the reserves of the stem cutting. • The most active vegetative growth for
• The fibrous roots start to grow, replacing cassava occurs in this period (Ramanujam,
the first adventitious roots. These new roots 1985).
start to penetrate in the soil, reaching
40–50 cm deep, and function in water and
High carbohydrate translocation to roots –
nutrient absorption (Conceição, 1979).
180–300 DAP
• Few fibrous roots (between three and 14)
will become storage roots, which can be • Photoassimilate partition from leaves to
distinguished from fibrous roots from 60 to roots is accelerated, making the bulking
90 DAP (Cock et al., 1979). At 75 DAP the of storage roots faster (Fig. 5.3).

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Botany and Physiology 75

• The highest rates of DM accumulation In cassava a positive correlation between the

in storage roots occur within this period leaf area or leaf area duration and yield of stor-
(Fig. 5.3; Boerboom, 1978; Távora et al., age roots has been reported, indicating that leaf
1995; Peressin et al., 1998). area is crucial in determining crop growth rate
• Leaf senescence increases, hastening rate of and the storage bulking rate of cassava (Sinha
leaf fall (Fig. 5.3). and Nair, 1971; Cock, 1976; Cock et al., 1979).
• Stem becomes lignified (Conceição, 1979). For cassava the leaf area per plant depends
on the number of active apices (branching
Dormancy – 300–360 DAP pattern), the number of leaves formed/apex, leaf
size and leaf life. Given that there are significant
• Rate of leaf production is decreased.
varietal variations and influence of environmen-
• Almost all leaves fall and shoot vegetative
tal conditions (Veltkamp, 1985a), it is important
growth is finished.
to characterize the development of cassava leaf
• Only translocation of starch to root is kept,
area and its components. Table 5.4 lists values
and maximum DM partition to the roots is
of some parameters related to leaf growth that
attained.
have been found in cassava. These parameters
• This phase occurs mainly in regions with
are discussed below.
significant variation in temperature and
After leaf emergence (folded, 1 cm long)
rainfall.
and under normal conditions, the cassava leaf
• The plant completes its 12-month cycle,
reaches its full size on days 10–12. Leaf life (from
which can be followed by a new period of
emergence to abscission) depends on cultivar,
vegetative growth, DM accumulation in the
shade level, water deficit and temperature (Cock
roots and dormancy again.
et al., 1979; Irikura et al., 1979). It ranges from
40 to 210 days (Table 5.4), but is commonly
60–120 days (Cock, 1984).
Leaf area development There are marked differences in leaf size
among the different cultivars, and the size varies
The analyses of crop growth and yield are with the age of the plant. The leaves produced
usually evaluated on the basis of two parame- from 3 to 4 MAP are those that become the larg-
ters: leaf area index (LAI), i.e. leaf area per unit est; maximum total leaf area is reached from 4 to
ground area, and net assimilatory rate (NAR), 5 MAP (Cock et al., 1979; Irikura et al., 1979).
i.e. the rate of DM production per unit leaf area. Leaf size is influenced by changing the branching

Table 5.4. Parameters related to leaf growth during the first cycle (12 months) and some values found
in cassava.

Leaf growth parameter Value Reference

Individual leaf area 50–600 cm2 Ramanujam (1982); Splittstoesser

and Tunya (1992); Veltkamp (1985a)
Expansion period (from emergence to 12 days Conceição (1979)
full size)
Leaf longevity (from fully expanded to 36–100 days Ramanujam and Indira (1983);
abscission) Conceição (1979)
Leaf life (from emergence to abscission) 40–210 days Splittstoesser and Tunya (1992);
Irikura et al. (1979); Ramanujam
(1985); Veltkamp (1985a)
No. leaves retained/plant 44–146 Ramanujam and Indira (1983)
Leaf production rate/shoot 4–22 week−1 Ramanujam and Indira (1983)
Leaf shedding rate/plant 10–24 leaves week−1 Ramanujam and Indira (1983)
Total leaf area 1.24–3.38 m2 Ramanujam and Indira (1983)
Cumulative no. of leaves/apex 117–162 Veltkamp (1985a); Cock et al. (1979)

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76 A.A.C. Alves

pattern. Larger leaves are produced when the

number of active apices is reduced (Tan and
Cock, 1979). The rate of leaf formation decreases
with plant age and is lower at low temperatures
(Irikura et al., 1979).
Differences in mean LAI are closely related
to the rate of root bulking. The optimal LAI for
storage root bulking rate is 3–3.5 (Cock et al.,
1979) and exists over a wide range of tempera-
ture (Irikura et al., 1979). Initial leaf area devel-
opment is slow, taking 60–80 DAP before an LAI
of 1.0 is reached. From 120 to 150 DAP the light
interception by the canopy is around 90% with
an LAI of 3 (Veltkamp, 1985a). In order to obtain
high storage root yields, the crop should reach an
LAI of 3–3.5 as quickly as possible and maintain Fig. 5.4. Temporal development of cell division
and cell expansion processes during cassava leaf
that LAI for as long as possible (Cock et al., 1979;
development in adaxial epidermal cells. (Source:
Veltkamp, 1985a). Substantial leaf abscission Alves, 1998.)
began at LAI values of 5.0–6.0 (Keating et al.,
1982a).
accumulation in the leaves decreases during the
crop cycle. Until 60–75 DAP, cassava accumu-
Temporal development of cell division lates DM more in leaves than in stems and
and cell expansion storage roots, not including the stem cutting
(Fig. 5.2). Then the storage roots increase
Leaf area development is largely dependent on rapidly, reaching 50–60% of the total DM
cell division and cell expansion processes, which around 120 DAP (Fig. 5.2; Howeler and
determine the number of cells per mature leaf Cadavid, 1983; Távora et al., 1995). After the
and cell size, respectively. Thus the final leaf area fourth month, more DM is accumulated in
is directly affected by the rate and duration of the storage roots than the rest of the plant. At
cell division and expansion (Takami et al., 1981; harvest (12 months) DM is present mainly in
Lecoeur et al., 1995). A temporal development roots, followed by stems and leaves (Fig. 5.3;
of cell division and expansion for adaxial epider- Howeler and Cadavid, 1983). Thus, during the
mal cells in cassava has been proposed by Alves growth cycle, the DM distribution to the differ-
(1998; Fig. 5.4), in which the transition from ent parts is constant with a high positive linear
leaf cell division to cell expansion processes is correlation of the total DM with shoot and root
discrete and occurs when leaf area reached 5% DM (Fig. 5.1; Veltkamp, 1985d).
of its final size, corresponding to the first folded The period of maximum rates of DM
leaf toward the top. Thus when the leaf starts to accumulation depends on genotypes and grow-
unfold, almost all cell division stops and rapid ing conditions. Lorenzi (1978) at high latitude
cell expansion starts. and Oelsligle (1975) at high altitude reported
maximum rates of DM accumulation at 4–6
and 7 months, respectively. Under more tropical
Dry matter partitioning and source–sink conditions, where growth is faster, Howeler and
relationship Cadavid (1983) found an earlier period of maxi-
mum rates, at 3–5 MAP. The distribution of DM
During cassava growth the carbohydrates from to the economically useful plant parts is mea-
photosynthesis have to be distributed to assure sured by harvest index (HI). In cassava HI repre-
good development of the source (active leaves) sents the efficiency of storage root production
and provide DM to the sink (storage roots, stem and is usually determined by the ratio of storage
and growing leaves). Cassava DM is translocated root weight to the total plant weight. Significant
mainly to stems and storage roots, and DM differences in HI have been reported among

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Botany and Physiology 77

cultivars, indicating that it can be used as a selec- (27° 37′ S), where photoperiods range from 14.8
tion criterion for higher yield potential in cas- to 11.2 h. Concentration of first flowering and
sava. HI values of 0.49–0.77 have been reported forking occurred in photoperiods > 13.5 h. This
after 10–12 MAP (Lorenzi, 1978; Cavalcanti, result is consistent with Bruijn (1977) and
1985; Pinho et al., 1995; Távora et al., 1995; Cunha and Conceição (1975), who suggested
Peressin et al., 1998). Although DM distribution flowering in cassava may be promoted by
is constant, its accumulation depends upon increasing day length.
photoassimilate availability (source activity) and
sink capacity of the storage parts. The number of
storage roots and their mean weight are yield
components that determine sink capacity. The Photosynthesis
significant positive correlation of photosynthetic
rate with root yield and total biomass, as well Cassava photosynthesis follows a C3 pathway
as the correlations between LAI, interception (Veltkamp, 1985e; Edwards et al., 1990; Angelov
of radiation and biomass production (Williams, et al., 1993; Ueno and Agarie, 1997) with
1972; Mahon et al., 1976; El-Sharkawy and maximum photosynthetic rates varying from
Cock, 1990; Ramanujam, 1990), indicates that 13 to 24 µmol CO2 m−2 s−1 under greenhouse
demand for photoassimilates by roots increases or growth chamber conditions (Mahon et al.,
the photosynthetic activity. 1977b; Edwards et al., 1990) and from 20 to
The balance between ‘source’ and ‘sink’ 35 µmol CO2 m−2 s−1 in the field (El-Sharkawy
activity is essential for the plant to reach its and Cock, 1990). It exhibits a high CO2 compen-
maximum productivity. Studies have shown sation point, from 49 to 68 µl l−1, typical of C3
that up to 25% reduction in the number of plants (Mahon et al., 1977a; Edwards et al.,
storage roots did not affect total or root DM and 1990; Angelov et al., 1993). In field-grown
the IAF (Cock et al., 1979). On the other hand, cassava, photosynthesis has high optimum
Ramanujam and Biradar (1987) observed that temperature (35°C) and wide plateau (25–35°C;
reduction of 50–75% in storage roots did affect El-Sharkawy and Cock, 1990) and is not light
root growth rate without changing shoot growth saturated up to 1800 µmol PAR m−2 s−1
rate, indicating that shoot growth is independent (El-Sharkawy et al., 1992b) or 2000 µmol PAR
of storage root growth. Influence of source size on m−2 s−1 (Angelov et al., 1993). Thus cassava
DM production shows that the NAR and storage is adapted to a tropical environment, requiring
root growth rate is reduced when the source size high temperature and high solar radiation for
is increased from LAI 3.0 to 6.0 (Ghosh et al., optimal leaf development and for expression of
1988). its photosynthetic potential. Both storage root
yield and total biomass show positive correlation
with photosynthesis rate (El-Sharkawy and
Flowering Cock, 1990; Ramanujam, 1990).
Morphologically, cassava leaves combine
Little is known about flowering in cassava, and some novel characteristics related to high
some clones have never been known to flower. productivity and drought tolerance and, conse-
Flowering can start 6 weeks after planting quently, to photosynthesis. The lower mesophyll
although the precise flowering time depends surface is populated with papillose-type epi-
on cultivar and environment. It appears that dermal cells, while the upper surface is fairly
cassava flowers best at moderate temperatures smooth, with scattered stomata and trichomes.
(approximately 24°C). It has been suggested The papillae appear to add about 15% to leaf
that forking is related to the onset of flowering, thickness and to lengthen the diffusion path
which is promoted by long days in some from the stomatal opening to the bulk air perhaps
cultivars. Usually, the apical meristem becomes two- to threefold (Angelov et al., 1993). Cassava
reproductive when branching occurs, but the leaves have distinct green bundle-sheath cells,
abortion of flowers is very common. with small, thin-walled cells, spatially separ-
Keating et al. (1982a) evaluated cassava ated below the palisade cells (different from
at 12 different planting dates at a high latitude Kranz-type leaf anatomy). In addition to

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78 A.A.C. Alves

performing C3 photosynthesis, these cells may fermentation, drying or a combination of these

function in transport of photosynthates in the processing treatments aid in reducing the HCN
leaf. In Kranz anatomy, typical of C4 plants, the concentration to safe levels (O’Hair, 1990).
bundle sheaths are surrounded by and all in
direct contact with many mesophyll cells
(Edwards et al., 1990). Physical Deterioration of
Storage Roots

Cyanide Content Cassava roots have the shortest postharvest

life of any of the major root crops (Ghosh et al.,
All cassava organs, except seeds, contain cyano- 1988). Roots are highly perishable and usually
genic glucoside (CG). Cultivars with < 100 mg become inedible within 24–72 h after harvest
kg−1 fresh weight (FW) are called ‘sweet’ while due to a rapid physiological deterioration pro-
cultivars with 100–500 mg kg−1 are ‘bitter’ cess, in which synthesis of simple phenolic com-
cassava (Wheatley et al., 1993). The most abun- pounds that polymerize occurs, forming blue,
dant CG is linamarin (85%), with lesser amounts brown and black pigments (condensed tannins;
of lotaustralin. Total CG concentration depends Wheatley and Chuzel, 1993). It is suggested that
on cultivar, environmental condition, cultural polyphenolic compounds in the roots oxidize to
practices and plant age (McMahon et al., 1995). quinone-type substances, which is complexed
The variation found in some parts of cassava is with small molecules like amino acids to form
shown in Table 5.5. coloured pigments that are deposited in the
Linamarin, which is synthesized in the leaf vascular bundles (Ghosh et al., 1988). The
and transported to the roots, is broken down by accumulation of the coumarin, scopoletin, is
the enzyme linamarase, also found in cassava especially rapid, reaching 80 mg kg−1 dry
tissues (Wheatley and Chuzel, 1993). When weight (DW) in 24 h (Wheatley and Chuzel,
linamarin is hydrolysed, it releases HCN, a vola- 1993). Tissue dehydration, especially at sites of
tile poison (LD50–60 mg for humans; Cooke and mechanical damage to the roots encourages the
Coursey, 1981); but some cyanide can be detoxi- rapid onset of deterioration. The phenolic com-
fied by the human body (Oke, 1983). In intact pounds may be released on injury. The changes
roots the compartmentalization of linamarase during storage of roots depend upon condition
in the cell wall and linamarin in cell vacuoles and duration of storage, physiological state of
prevents the formation of free cyanide. Upon the stored material and varietal characteristics
processing, the disruption of tissues ensures that (Ghosh et al., 1988).
the enzyme comes into contact with its substrate, Polyphenol oxidase (PPO) is an enzyme
resulting in rapid production of free cyanide via that oxidizes phenols to quinone. Any process
an unstable cyanhydrin intermediary (Wheatley that inhibits PPO, such as heat treatment, cold
and Chuzel, 1993). Juice extraction, heating, storage, anaerobic atmosphere and dipping

Table 5.5. Cyanide concentration in different parts of the cassava plant.

Part of plant Total cyanide concentration Source

Root pulp (parenchyma) 3–121 mg 100 g−1 DW Barrios and Bressani (1967)
3–135 mg 100 g−1 DW Wheatley and Chuzel (1993)
1–40 mg 100 g−1 FW Barrios and Bressani (1967)
Root peel 6–55 mg 100 g−1 DW Wheatley and Chuzel (1993)
5–77 mg 100 g−1 DW Barrios and Bressani (1967)
17–267 mg 100 g−1 FW Barrios and Bressani (1967)
Leaf 1–94 mg 100 g−1 DW Barrios and Bressani (1967)
0.3–29 mg 100 g−1 FW Barrios and Bressani (1967)

DW, dry weight; FW, fresh weight.

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Botany and Physiology 79

roots in solutions of inhibitors (e.g. ascorbic of cassava under the temperature variations
acid, glutathione and KCN) prevents vascular that usually occur where cassava is normally
streaking (Ghosh et al., 1988). cultivated indicates that its growth is favourable
Secondary deterioration can follow physio- under annual mean temperatures ranging from
logical or primary deterioration 5–7 days after 25 to 29°C (Conceição, 1979), but it can
harvest. This is due to microbial infection of tolerate from 16 to 38°C (Cock, 1984). Table
mechanically damaged tissues and results in 5.6 summarizes the ranges of temperature and
the same tissue discoloration with vascular their principal physiological effects on cassava
streaks spreading from the infected tissues development.
(Wheatley and Chuzel, 1993). At low temperatures (16°C) sprouting of
the stem cutting is delayed, and rate of leaf
production, total and storage root DW are
Environmental Effects on decreased (Cock and Rosas, 1975). Sprouting
Cassava Physiology is hastened when the temperature increases
up to 30°C but is inhibited with temperatures
Cassava is found over a wide range of edaphic > 37°C (Keating and Evenson, 1979). As
and climatic conditions between 30°N and 30°S temperature decreases, leaf area development
latitude, growing in regions from sea level to becomes slower because the maximum size of
2300 m altitude, mostly in areas considered individual leaves is smaller, and fewer leaves
marginal for other crops: low-fertility soils, are produced at each apex although leaf life is
annual rainfall from < 600 mm in the semiarid increased (Irikura et al., 1979). At a temperature
tropics to > 1500 mm in the subhumid and of 15–24°C, the leaves remain on the plant for up
humid tropics. Given the wide ecological to 200 days (Irikura et al., 1979), while at higher
diversity, cassava is subjected to highly varying temperatures leaf life is 120 days (Splittstoesser
temperatures, photoperiods, solar radiation and and Tunya, 1992).
rainfall. There is a genotype-by-temperature inter-
action for yield ability. Irikura et al. (1979) evalu-
ated four cultivars under different temperatures
Temperature and found that higher yields were obtained at
different temperatures according to the cultivar,
Temperature affects sprouting, leaf size, indicating that the effect of natural selection
leaf formation, storage root formation and, con- is highly significant on varietal adaptation
sequently, general plant growth. The behaviour (Table 5.7).

Table 5.6. Effect of temperature on cassava development.

Air temperature (°C) Physiological effects

< 17 or > 37 Sprouting impaired

28.5–30 Sprouting faster (optimum)
< 15 Plant growth inhibited
16–38 Cassava plant can grow
25–29 Optimum for plant growth
< 17 Reduction of leaf production rate, total and root DW
20–24 Leaf size and leaf production rate increased; leaf life shortened
28 Faster shedding of leaves; reduction in no. branches
25–30 Highest rates of photosynthesis in greenhouse
30–40 Highest rates of photosynthesis in the field
16–30 Transpiration rate increases linearly and then declines

Sources: Wholey and Cock (1974); Cock and Rosas (1975); Mahon et al. (1977b); Conceição (1979);
Irikura et al. (1979); Keating and Evenson (1979); El-Sharkawy et al. (1992b).
DW, dry weight.

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80 A.A.C. Alves

Table 5.7. Fresh root yield (t ha−1) of four contrasting cassava types at 12
months after planting (MAP) under three different temperature regimes.

Temperature (°C)

Variety 20 24 28

M Col 22 9.3 27.7 39.4

M Mex 59 22.8 38.8 30.4
M Col 113 24.2 26.1 23.9
Popayán 28.9 15.7 9.4

Source: Irikura et al. (1979).

The main effect of temperature is on bio- length (Bolhuis, 1966). Long days promote
logical production, as DM partitioning does not growth of shoots and decrease storage root
change much when cassava is cultivated under development, while short days increase storage
different temperatures (Cock and Rosas, 1975). root growth and reduce the shoots, without
Higher temperatures are associated with a influencing total DW (Fig. 5.5). The increase
greater crop growth rate (CGR) and high photo- in shoot DW under long days is a result of
synthetic rate. El-Sharkawy et al. (1992b) significant increases in plant height, leaf area
evaluated the potential photosynthesis of three per plant, number of apices per plant, and
cultivars from contrasting habits under different number of living leaves per apex (Veltkamp,
growing environments and verified that photo- 1985b). This suggests an antagonistic relation-
synthetic rate increased with increasing temper- ship between shoot growth and development of
ature, reaching its maximum at 30–40°C. In all the storage roots in response to variation in day
cultivars photosynthesis was substantially lower length.
in leaves that had developed in the cool climate There are varietal differences in sensitivity
than in those from the warm climate. The high to long days (Carvalho and Ezeta, 1983). Under
sensitivity of photosynthesis to temperature field conditions, Veltkamp (1985b) submitted
points to the need for genotypes more tolerant three genotypes to two day lengths (12 and 16 h)
to low temperature, which could be used in the during the whole growth period. He observed
highland tropics and subtropics. that the percentage decrease in storage root yield
under the long day was greatest in M Col 1684
(47%) and least in M Col 22 (13%; Table 5.8).
Photoperiod Yield differences resulted mainly from decreased
efficiency of storage root production or HI under
Day length affects several physiological pro- 16-h days because total DM was greater under
cesses in plants. The differences in day length 16-h days. No day-length effect was found for the
in the tropical region are very small, varying crop weight at which the starch accumulation in
from 10 to 12 h throughout the year. Thus the roots apparently started (AISS values; Table
photoperiod may not limit cassava root pro- 5.8). Thus the storage root yield reduction seems
duction in this region. On the other hand, the to be more related to a change in the distribution
restrictions regarding cassava distribution out- patterns of DM rather than to a delay in storage
side the tropical zone can be due to effects of day root initiation. Considering that photoperiod pri-
length variation on its physiology. Although marily affects shoots and a secondary response
studies about day length effect in cassava are occurs in the roots (Keating et al., 1982b) and
scarce, tuberization, photoassimilates partition- that shoot growth has preference over root
ing and flowering are reportedly affected. growth (Cock et al., 1979; Tan and Cock, 1979),
Experiments in which the day length was long photoperiods may increase the growth
artificially changed have shown that the optimal requirements of the shoots, thereby reducing the
light period for cassava is around 12 h, with excess carbohydrates available for root growth
probable varietal differences in the critical day (Veltkamp, 1985b).

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Botany and Physiology 81

Fig. 5.5. Effect of day length on cassava dry matter distribution 16 weeks after planting. Vertical lines
indicate twice the standard error of total dry weight (DW), stem DW and storage root DW.

Table 5.8. Cassava dry matter (DM) production and distribution 272 days after planting (DAP) under
16 h and natural day length (approximately 12 h).

Total DM DW storage root Harvest Index AISSb

Genotype Day length (t ha−1)a (t ha−1) (HI) (t ha−1)

M Col 1684 Naturalc 16.7 9.1 0.54 0.55

16 hd 17.3 4.6 0.27 0.25
M Ptr 26 Natural 14.5 8.1 0.61 0.60
16 h 15.9 4.9 0.42 0.36
M Col 22 Natural 15.5 9.5 0.56 0.65
16 h 19.5 8.3 0.31 0.45

Source: Veltkamp (1985b).

aIncludes weight of fallen leaves.
bAISS, apparent initial start of starch accumulation; corresponds to crop weight at which starch

accumulation apparently starts in roots.

cNatural = approximately 12 h.
d16 h = 12 h of natural day length + 4 h of artificial light.

DW, dry weight.

Solar radiation groundnuts; Mutsaers et al., 1993). Cassava

is also intercropped with perennial vegetation
The commonest cassava production system is (Ramanujam et al., 1984). Cassava is usually
intercropping with other staple crops. In Latin planted after the intercropped species. Even
America and Africa, cassava is usually associ- when it is planted at the same time, the associ-
ated with an earlier maturing grain crop such as ated crop such as maize is established faster than
maize, rice or grain legumes (beans, cowpeas or cassava. Thus in an associated cropping system

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82 A.A.C. Alves

cassava is always subjected to different degrees but under 95–100% shade, leaves abscise within
of shading and low light intensity in the early 10 days (Cock et al., 1979).
stages of development. Considering that cassava Aresta and Fukai (1984) observed that only
is a crop that requires high solar radiation 22% shade decreased both fibrous root elonga-
to perform photosynthesis more efficiently (El- tion rate (53%) and storage root growth rate
Sharkawy et al., 1992b), it is very important to (36%) without altering shoot growth rate, which
know the effect of shade on cassava develop- was significantly decreased (32%) only under
ment and production. Ramanujam et al. (1984) 68% shade. Thus under limited photosynthesis
evaluated 12 cassava cultivars under the shade caused by low solar radiation, most of the photo-
in a coconut garden (85–90% shading). Under synthates are utilized for shoot growth, affecting
shading, the root bulking process started about storage root development significantly, showing
3 weeks after that in plants grown without that the shoots are a stronger sink than roots.
shading, and the number of storage roots per
plant and NAR was reduced under shading.
Okoli and Wilson (1986) submitted cassava to Water deficit
six shade regimes and observed that all levels of
shade delayed storage root bulking and at 20, Cassava is commonly grown in areas receiving
40, 50, 60 and 70% shade reduced cassava yield < 800 mm rainfall year−1 with a dry season of
by 43, 56, 59, 69 and 80%, respectively. 4–6 months, where tolerance to water deficit is
In relation to shoots, under field conditions, an important attribute. Although it is a drought-
shading increases plant height and the leaves tolerant crop, growth and yield are decreased by
tend to become adapted to low light conditions by prolonged dry periods. The reduction in storage
increasing leaf area per unit weight (Fukai et al., root yield depends on the duration of the water
1984; Okoli and Wilson, 1986; Ramanujam deficit and is determined by the sensitivity of
et al., 1984) and shortening leaf life only under a particular growth stage to water stress. The
severe shading. Under ideal growing conditions, critical period for water-deficit effect in cassava
cassava leaves have a life of up to 125 days is from 1 to 5 MAP – the stages of root initiation
(Splittstoesser and Tunya, 1992). Levels of shade and tuberization. Water deficit during at least 2
up to about 75% have very little effect on leaf life, months of this period can reduce storage root

Fig. 5.6. Effect of water deficit during different growth periods on cassava yield.

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Botany and Physiology 83

yield from 32 to 60% (Connor et al., 1981; Porto 1984; Cock et al., 1985). This response to early
et al., 1988). Figure 5.6 shows the root yield stages of soil water depletion has been described
reduction caused by water deficit imposed for 2 as isohydric, a behaviour shared by cowpeas,
months during successive 2-month periods from maize and several other crops (Tardieu and
1 to 11 MAP. Clearly, they found that the Simonneau, 1998). Leaf area growth is also
severer effect corresponded to stress occurring decreased in response to water stress but is
from 1 to 5 MAP (i.e. period of rapid leaf growth rapidly reversed following the release from
and tuberization) compared with the later stress (Connor et al., 1981; Palta, 1984;
period of storage root bulking. El-Sharkawy and Cock, 1987b; Baker et al.,
1989). This response limits the development of
plant transpirational surface area during water
Drought tolerance deficit and keeps sink demand well balanced with
plant assimilatory capacity.
Plants respond to water deficit at many differ- Leaf conductance to water vapour has been
ent levels: morphological, physiological, cellular evaluated as an indicator of the capacity of
and metabolic. The responses are dependent different cassava genotypes to prevent water
upon the duration and severity of stress, the loss under prolonged drought. Considerable
genotype of the stressed plant, the stage of variation has been observed in leaf conductance
development, and the organ and cell type in (Porto et al., 1988), and this parameter seems
question (Bray, 1994). Multiple responses allow to be useful for pre-selecting sources of germ-
the plant to tolerate water stress. Some of these plasm conferring adaptation to prolonged dry
responses and the current status of knowledge periods.
with regard to cassava are discussed here.

Control of stomatal closure and Abscisic acid accumulation

leaf growth
Many authors have reported that a substantial
A primary response to water stress is stomatal number of drought responses in plants can
closure, which decreases photosynthetic CO2 be mimicked by external application of abscisic
assimilation and, in turn, growth. Stomata have acid (ABA) to well-watered plants (Davies et al.,
a high capacity to respond to changes in the 1986; Trewavas and Jones, 1991). This treat-
water status of the plant and atmosphere. They ment promotes characteristic developmental
close as the leaf water potential decreases and changes that can help the plant cope with water
when the vapour pressure deficit between the deficit, including decrease of stomatal con-
leaf and the air increases (generally due to a ductance (MacRobbie, 1991; Trejo et al., 1993),
decrease in relative humidity). As stomata are restriction of shoot growth (Creelman et al.,
the route by which CO2 enters the leaf, stress- 1990) and leaf area expansion (Van Volken-
induced decreases in stomatal aperture can burgh and Davies, 1983; Lecoeur et al., 1995)
limit the rate of CO2 diffusion into the leaf and, and stimulation of root extension (Sharp et al.,
therefore, the rate of photosynthesis. 1993, 1994; Griffiths et al., 1997). All these
When water is available, cassava maintains effects of ABA application, together with the
a high stomatal conductance and can keep observation that environmental stress stimu-
internal CO2 concentration high; but when lates ABA biosynthesis and ABA release from
water becomes limiting, the plant closes stomata sites of synthesis to action sites, suggest a role
in response to even small decreases in soil water for ABA as a stress hormone in plants. More-
potential (El-Sharkawy and Cock, 1984). The over, studies have indicated that for certain
rapid closure of the stomata and the resulting plant responses, sensitivity to water deficit is
decline in transpiration lessens the decrease correlated with changes in ABA concentrations
in leaf water potential and soil water depletion, (Trejo et al., 1995; Borel et al., 1997) and geno-
thereby protecting leaf tissues from desiccation typic responsiveness to ABA (Blum and Sinmena,
(Ike, 1982; Palta, 1984; El-Sharkawy and Cock, 1995; Cellier et al., 1998). Information about

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84 A.A.C. Alves

ABA accumulation in cassava has not been Osmotic adjustment

reported in the literature.
Alves and Setter (2000) published the One means of increasing drought tolerance is
first report concerning ABA in cassava. They by accumulation of osmotically active solutes,
cultivated five cassava genotypes in pots in a so that turgor and turgor-dependent processes
greenhouse and evaluated the temporal patterns may be maintained during episodes of drydown.
of ABA accumulation in mature leaves and in Osmotic adjustment (OA), defined as the differ-
immature expanding leaves, during water deficit ence in osmotic potential between control and
and after release from stress to determine the stressed plants, allows cell enlargement and
extent to which the stress and recovery response plant growth at high water stress and allows
of leaf area growth is associated with the tempo- stomata to remain partially open and CO2 assim-
ral pattern of ABA accumulation. At 3 and 6 ilation to continue at low water potentials that
days after withholding irrigation, all genotypes are otherwise inhibitory (Pugnaire et al., 1994).
accumulated large amounts of ABA in both In cassava OA has not been thoroughly
expanding and mature leaves, but these high examined. Although leaf water potential
ABA levels were almost completely reversed in remains relatively unchanged during water
respect of control levels after 1 day of rewatering deficit episodes (Connor and Palta, 1981; Cock
(Fig. 5.7). Correspondingly, young leaves halted et al., 1985; El-Sharkawy et al., 1992a), a result
leaf expansion growth and transpiration rate suggestive of little or no OA, such observations
decreased. Young leaves accumulated more do not rule it out. Furthermore, these studies
ABA than mature leaves both in control and involved mature leaves, not young organs,
stressed treatments (Fig. 5.7). This rapid return which might especially benefit from accumula-
to control ABA levels corresponded with a rapid tion of osmolytes. Alves (1998) evaluated the
recovery of leaf area growth rates. This rapid role of OA in cassava by measuring the osmotic
reduction in leaf area growth and stomatal component of leaf water potential in mature,
closure may be due to cassava’s ability to expanding and folded leaves. The largest
synthesize rapidly and accumulate ABA at an increase in solutes caused by water deficit
early phase of a water-deficit episode. occurred in the youngest tissue (folded leaves)

Fig. 5.7. Abscisic acid (ABA) concentration in expanding and mature cassava leaves in control after 3
and 6 days of water deficit, followed by 1 and 3 days of rewatering; average of five genotypes with three
replicates; bars represent SEM (n = 15); data from Alves and Setter (2000).

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Botany and Physiology 85

and the extent of OA increased progressively Borel, C., Simonneau, T., This, D. and Tardieu, F.
from mature to folded leaves. As young tissues (1997) Stomatal conductance and ABA con-
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