Sunflower Physiology

Published 1997
4 Sunflower Physiology
DAVID J. CONNOR
University of Melbourne
Parkville 3052
Australia
ANTHONY J. HALL
University of Buenos Aires
Buenos Aires, Argentina
Physiology is concerned with the development, growth, and yield of plants in

response to environmental conditions. It focuses on several levels of organiza-
tion, from subcellular to the community (crop), bridging the gap between bio-
chemistry and agronomy. Just as agronomists seek, in physiology, explanations
for the responses of crops they observe in the field, so too, physiologists in tum,
seek explanations of the integrated behavior of plants in the biochemistry of
metabolism.
This chapter treats the physiology of sunflower (Helianthus annuus L.)
with outward glances to agronomy and inward glances to biochemistry. The
sequence of topics reflects the nature of growth and the development of yield in
any crop. The water, nutrient, and C economies determine growth, but the distri-
bution of biomass to the various plant organs, and the nature of crop-environment
interactions for a given crop, depend upon phenological development, so that
topic is treated first. The water, nutrient, and C balances of sunflower have been
well studied in recent years and much information is now available. The high
growth rates and drought tolerance of sunflower have provided considerable
impetus to researchers to improve both those aspects. In contrast to the produc-
tive processes themselves, the controls over the partitioning of growth to compo-
nent organs, and in particular to seed, are much less certain. However, there is the
opportunity to make comparisons with other crops to propose possible explana-
tions for the behavior of sunflower, and in so doing to identify opportunities for
research. The economic yield of sunflower lies in the oil and protein content of
its seed, so there is a need to explore the metabolic interconversions by which the
end-products of photosynthesis are elaborated.
While it is necessary to deal with processes singly and present detail piece
by piece, it is the objective of physiology to explain how they interact to deter-
mine crop performance. For that reason, the chapter concludes with two views of
the integration of physiological responses. The first is a simple one that looks at
Copyright rg 1997. American Society of Agronomy, Crop Science Society of America, Soil Science
Society of America, 677 S. Segoe Rd., Madison, WI 53711, USA. Sunflower Technology and Produc-
tion, Agronomy Monograph no. 35.
113
114 CONNOR & HALL
pairs of important responses (e.g., growth vs. radiation capture, growth vs. water
use) and considers how linkages can be strengthened or modified by genotype or
management. The second introduces simulation modeling as a way to study the
interaction of the many chemical and physical processes that comprise plant
development, growth, and the expression of yield. There has been significant
progress in the application of these techniques to the study of sunflower and to
the integration of knowledge to assist plant improvement and crop management.
PHENOLOGICAL DEVELOPMENT
The time elapsed between sowing and harvesting of sunflower crops is
under strong environmental control, and there is significant variation in response
between genotypes. This is particularly so for the period from sowing to flower-
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Emergence day (after Dec. 1)
Fig. 4-1. (a) Duration of emergence to anthesis to anthesis for short- (SH 3000), medium- (894), and
long-cycle (Sungro 380) sunflower hybrids sown at different times during a single growing season.
(b) Duration of emergence to anthesis for SH 3000 sown at different times in four growing seasons
[data from experiments by Gimeno et a!. (1989); figures from Villalobos et a!. (1994)].
SUNFLOWER PHYSIOLOGY 115
ing, which responds markedly to date of sowing and interannual variations in

environmental conditions (Fig. 4-1). The understanding and, where possible, the
quantitative description of these genotype/environment interactions is a powerful
aid to agronomists and breeders in their quest to minimize the effects of stress and
to achieve the best match between yield potential and environment (e.g., Goyne
et aI., 1978; Beard & Geng, 1982; Gimeno et aI., 1989; Sadras & Villalobos,
1994). Related issues are the effects of environment, cultivar, and storage condi-
tions on seed dormancy, its breakage, and on seed aging. Dormant seed will not
germinate until dormancy is broken and seed stored under inappropriate condi-
tions lose vigor and germinability.
The phenological development of crops between sowing and physiological
maturity is conveniently considered as a sequence of phenophases separated by
identifiable phenostages. The environmental factors controlling the duration of
each phase vary in identity and relative importance. This account concentrates on
the phases from sowing to emergence (S-E), emergence to floral initiation (E-FI),
floral initiation to first anthesis (FI-FA), first anthesis to last anthesis (FA-LA),
and last anthesis to physiological maturity (LA-PM). In addition to describing the
phases and identifying the environmental factors that affect them, we also review
the physiology of the underlying processes and attempt to quantity development
in response to environment.
Clear identification of phenostages is essential, and several scales based on
visible changes in crop morphology and color have been developed (e.g.,
Siddiqui et aI., 1975; Anderson et aI., 1978; Schneiter & Miller, 1981). However,
identification of FI and monitoring the early development of the inflorescence
require dissection for which Marc and Palmer (1981) have devised a useful scale.
Recently, Sadras and Villalobos (1993) have shown how to establish the timing
of FI, at least for unstressed crops, from the progress of leaf emergence and the
final number of leaves. There is not yet, however, a simple technique for identi-
fication of PM, although changes in head color and ease of perianth abscission
have been proposed (Anderson et aI., 1978; Schneiter & Miller, 1981; Robinson,
1983; Browne, 1978). These subjective techniques serve to rank genotypes for
maturity in individual environments, but for broader comparisons, and for
research, the tedious technique of following seed or embryo dry-weight dynam-
ics to determine the timing of maximum weights is essential (e.g., Anderson,
1975; Hall et aI., 1985).
Quantitative analyses of crop development rely heavily on the concepts of
thermal time and formalized descriptions of photoperiodic responses. Ritchie and
NeSmith (1991), Loomis and Connor (1992), Roberts et aI. (1993), and Slafer
and Rawson (1994) provide useful discussions of these concepts and identify
some of the pitfalls associated with their use.
Seed Dormancy and Aging
Sunflower embryos develop the capacity for germination about 6 dafter

pollination and then become dormant about 10 d later (16 d after pollination),
which is close to the time when they develop tolerance to desiccation, but well
before the accumulation of seed reserves is complete. Abscisic acid (ABA)
116 CONNOR & HALL
appears to be involved in the induction of both desiccation tolerance and dor-

mancy, and changes in embryo sensitivity to ABA may affect the establishment
of dormancy (Le Page-Degivry et aI., 1990, 1992; Le Page-Degivry & Garello,
1991). The existence of a period of transient germinability soon after pollination
makes embryo rescue a feasible technique in sunflower breeding and biotechnol-
ogy (Alissa et aI., 1986). Some loss of embryo dormancy may occur in the final
stages of maturation (Corbineau et aI., 1990) but sunflower seed may exhibit
marked dormancy at PM. Low germination percentages attributable to dormancy
are seen most clearly at temperatures lower than the optimum (Corbineau et aI.,
1990). By analogy with other species, the degree of dormancy present at harvest
may vary with the conditions experienced during seed formation (Fenner, 1991).
Le Page-Degivry et al. (1990) reported year-to-year differences in the degree of
embryo dormancy, while Wallace and Habermann (1958) reported differences
between cultivars. Embryo dormancy, that is lost during several months of dry
storage depending upon conditions, is seen in increasing germination percentages
and gradual relaxation of temperature requirements for germination.
The seed envelope (pericarp and true seed coat) also contributes to dor-
mancy, being especially important at temperatures above 25 DC. Exposure of seed
to ethylene and its precursors can hasten breakage of dormancy of whole seed,
embryos enclosed in seed coats, and isolated embryos (Corbineau et aI., 1990).
Periods from 2 to more than 8 wk have been reported for breakage of whole seed
dormancy (Zimmerman & Zimmer, 1978; Maeda & Ungaro, 1985; Corbineau et
aI., 1990), but in these reports different criteria were used to define dormancy
breakage. In addition to ethylene, wet scarification, rinsing with water, exposure
to high concentrations of CO 2 or 02, and alternating temperatures during incuba-
tion, can hasten breakage of dormancy (Harada, 1982; Maeda & Ungaro, 1985;
Corbineau et aI., 1990).
Theory supports the proposition that there is an optimum seed moisture
content for prolonged storage and evidence suggests that this may be close to 3%
dry-weight for sunflower (Vertucci & Roos, 1990, 1993). This value is lower than
for many other species, consistent with the high oil content of the embryo.
Although studies of the effects of storage conditions on aging and the loss of
vigor and viability in sunflower have been limited to seed moisture content
(Vertucci & Roos, 1990), these processes should, by analogy with other species,
be hastened by high temperature and supraoptimal moisture content (Ellis &
Roberts, 1980). Seed quality can be an important factor affecting establishment,
and seedling emergence correlates better with variables that reflect seed condition
(leachate conductivity and germination at low temperature or after accelerated
aging) than with a standard germination test at 21 DC (Anfinrud & Schneiter,
1984).
Sowing to Emergence
This phase includes two distinct subphases, germination (start of imbibition

to protrusion of the radicle) and emergence (radicle protrusion to the unfolding of
the hypocotyledonary hook above the soil surface) (Bewley & Black, 1985).
Agronomic experiments rarely distinguish between these subphases so interpre-
tation of results is often difficult. Early stages in germination (radicle elongation)

reflect expansion of existing cells rather than cell division and the synthesis of the
requisite enzymes is linked to translation rather than transcription (Lenormand et
aI., 1993). Continued growth of seedlings presumably reflects further changes in
seed physiology and biochemistry such as oil-body breakdown (Bewley & Black,
1985; Murphy, 1990; Huang, 1992), but this is unexplored in sunflower.
Temperature is the most important factor affecting the germination of sun-
flower in soils with adequate water and aeration. Estimates ofthe minimum tem-
perature for germination lie in the range 3 to 6°C, optimum temperatures are
close to 26°C, and maximum temperatures approximately 40°C (Roilier et aI.,
1977, p. 3-28; Cseresnyes, 1979; Macchia et aI., 1985; Maeda & Ungaro, 1985;
Gay et aI., 1991). Oil composition (Downes, 1985) may affect germination rate,
particularly at low temperatures. Field establishment, measured 8 wk after a win-
ter sowing, increased from 10 to 95% as linoleic acid concentration increased
from 530 to 730 g/kg fatty acid (Downes, 1985). Temperature also affects emer-
gence of germinated seeds which is delayed about l2°Cd (degree days, summa-
tion of daily mean temperature above a base temperature, Tb, here = 4°C) per cen-
timeter sowing depth (Villalobos et aI., 1994). Comparative studies across geno-
types of the temperature responses of germination and emergence are urgently
needed and could provide important insights into intraspecific variability of these
processes, as was shown for maize (Zea mays L.) by Eagles and Hardacre (1979).
Under laboratory conditions, sunflower germination was restricted at water
potentials between -1.1 and -1.5 MPa, and was completely inhibited below -2.1
MPa (Somers et aI., 1983), while Chimenti (1991) found that water potential
thresholds for 50% germination varied between -0.7 and -2.2 MPa across a
range of genotypes. In a silty loam soil, germination and emergence proved even
more sensitive, with thresholds increasing to -0.7 and -1.4 MPa, respectively, for
restriction and complete inhibition (Somers et aI., 1983). Those authors also
found intraspecific variability for tolerance to low water potential. Sunflower ger-
mination is sensitive to O2 partial pressure (Gay et aI., 1991) with obvious, but as
yet unexplored, implications for sensitivity to waterlogging in field conditions.
There appears to be some intraspecific variability in tolerance of germination,
hypocotyl elongation, and emergence to salinity (Karami, 1974; Hardwick &
Ferguson, 1978). The emergence of seedlings derived from large seeds may
sometimes (Radford, 1977; Hocking & Steer, 1989), but not always (Cholaky et
aI., 1982), occur faster or from greater depths than for small seeds. General con-
clusions cannot be drawn from these results because seedbed characteristics
(crumb ·size, water content) interact with seed size (area/volume ratio) and sow-
ing depth (Bewley & Black, 1985). More precise studies are needed to consider
individual aspects of the response and possible cultivar effects such as slower
rates of hypocotyl elongation in dwarf cultivars (Sadras, 1994, personal commu-
nication).
Deficiencies in germination and/or emergence result in uneven stands, so
seed quality, uniformity of seed placement during sowing, and seedbed condi-
tions have important effects on stand quality. Further issues with sunflower are
the irregular shape of the seed and the small size in modern hybrids compared
with older, open-pollinated cultivars. Seed coating (Allen et aI., 1983) can
118 CONNOR & HALL
increase size and make shape more uniform to facilitate sowing, as well as
increasing germination rates. Priming of seeds by soaking in water or various
solutions followed by drying before sowing also can improve germination
(Kathiresan & Gnanarethinam, 1985; Smok et aI., 1992). Several possible reasons
for this response in other species have been advanced (Bewley & Black, 1985;
Dahal et aI., 1990), but its physiological basis remains unknown. Although prim-
ing is unlikely to be important for extensive sowings, it may be a useful technique
for dealing with small samples oflow-vigor seed in germplasm-maintenance pro-
grams. The potentially damaging consequences of uneven stands in sunflower is
compensated to some extent by the plasticity of leaf and root growth (Sadras et
aI., 1989). Cardinali et ai. (1985) found no differences in yield over a range of
crop density from 1.8 to 7.2 plants/m2 between homogeneous stands and crops in
which the emergence of 50% of the population had been delayed by 1 wk.
Angus et ai. (1981) established a quantitative description of the effect of
temperature on the duration of germination to establishment using data from
time-of-sowing experiments. Villalobos et ai. (1994) extended this in controlled-
environment experiments with seed sown at various depths to distinguish
between germination and emergence. However, more detailed studies of the tem-
perature response of both processes are still needed, using nondormant seed, with
appropriate identification of base, optimal, and maximum temperatures (Garcia-
Huidobro et aI., 1982). These studies also must investigate the intraspecific vari-
ability in temperature sensitivity associated with variations in linoleic acid
(C 1s H320 2) content (Downes, 1985). Extension of these efforts to encompass
other important factors such as soil water content and seedbed characteristics also
is needed. An issue of particular interest is the supposedly greater sensitivity of
sunflower germination and emergence to waterlogging, compared with cereals.
Emergence to Floral Initiation
The duration ofE to FI comprised approximately 20% of the growth cycle

of ' Suncross 150' grown under high irradiance and a range of constant tempera-
tures (Rawson et aI., 1984) and between 20 and 27% of the thermal duration of
four hybrids of varying maturity grown in the field (Villalobos et aI., 1994).
Floral initiation signals the end of the production of leaf primordia by the shoot
apex and the beginning of reproductive activity. It is, therefore, an important
event in crop development that warrants its tedious identification by dissection of
the apex.
There are several independent indications, although no experimental evi-
dence, that sunflower has a juvenile phase (Vince-Prue, 1975), i.e., a "condition"
that must be overcome before development can proceed to FI. Changes in mitot-
ic activity of the central portion of the shoot apical meristem, the first micro-
scopically detectable indication of the changes that precede FI (Steeves et aI.,
1969; Langenauer et aI., 1975), occurred around 8 d after sowing in sunflower
seedlings grown at (the fairly high) temperature of28°C (Marc & Palmer, 1982).
Time to visible differentiation of the inflorescence was shortened by grafting
seedling apical meristems onto stocks more than 40 d old, and this effect
increased with age of the stock (Habermann & Wallace, 1958). Manipulation of
development by exposing seedlings to short chilling pulses (Marc & Palmer,

1978) revealed the existence of a temperature-insensitive phase 8 d after sowing
or about 10 d before visible differentiation of the floral apex. Although FI and
leaf primordia formation can be shown to be partially independent under stress
(Marc & Palmer, 1976, 1978), they are closely related in unstressed plants
(Sadras & Villalobos, 1993). The results of Marc and Palmer (1978) suggest that
a substantial number of leaf primordia, more than the two present in the embryo
(von Guttenberg et aI. , 1955), were initiated before the start of the temperature-
insensitive phase.
The duration of E to FI in sunflower can, therefore, be expected to depend
on the duration of the juvenile phase plus the time needed for the changes in the
apical meristem to become visible. By analogy with other species (e.g., maize,
Kiniry et aI., 1983), photoperiodically sensitive cultivars of sunflower should not
respond to photoperiod until the juvenile phase is complete, when exposure to a
minimum number of inductive cycles may be required to evoke FI (Vince-Prue,
1975). This hypothesis has not been tested experimentally.
Marc and Palmer (1981) developed a scale to describe the progress of inflo-
rescence development between the first changes in shape of the meristem [their
floral stage (FS) I] and the completion of gross differentiation of the florets (FS
10, all central florets with visible corolla and hairy floret bracts). Important stages
in this scale are FS 1.3 (i.e., the stage that coincides with the differentiation of the
last leaf primordia, and used in this review to indicate floral initiation); FS 5 when
the primordia of the ray florets appear at the periphery of the floral disc; and the
end of FS 7 to the start of FS 8, when floret differentiation in the central portion
of the receptacle is complete.
The duration of E to FI for various cultivars, when examined in controlled
environment conditions, using reasonable irradiance levels and protocols that
avoid confounding the effects of photosynthesis and photoperiod, was shown to
be differentially sensitive to temperature and photoperiod (Rawson &
Hindmarsh, 1982; Rawson et aI., 1984). These responses have been confirmed
(Goyne & Hammer, 1982; Goyne & Schneiter, 1988) in experiments in which
bud visible (BV) (extemally visible inflorescence, R 1 in Schneiter and Miller
(1981), and approximately FS 7 in Marc and Palmer (1981)) was used as a sur-
rogate for FI. Substantial differences between cultivars for the duration E to FI
also have been found in field experiments (Villalobos et aI., 1994).
Most sunflower cultivars exhibit quantitative long-day or day-neutral
responses to photoperiod for the duration of E to FI (or E-BV), but at least one
cultivar has exhibited a short-day response (Rawson & Hindmarsh, 1982; Goyne
& Schneiter, 1988). There has been considerable discussion about the nature of
photoperiodic responses in sunflower and this is considered later in this review.
Here we note that caution should be exercised in accepting the classifications
reported in early literature, when FI was not identified and experimental condi-
tions were not always appropriate for correct categorization of the photoperiodic
response. Extrapolation from other species (Major, 1980) suggests that the pho-
toperiodic responses of sunflower should exhibit threshold and (possibly) critical
values, i.e., for a long-day response, the minimum and maximum photoperiods
that affect developmental rate. In one set of experiments these values were esti-
120 CONNOR & HALL
mated as 13 and 15 h (Hammer et aI., 1982); in another, all photoperiods includ-

ed in the experimental range (10-15 h) affected the rate of development (Rawson
& Hindmarsh, 1982).
While temperature and photoperiod are clearly the important factors affect-
ing the duration of E to FI, other effects also have been recorded. Reduction of
irradiance from 25.5 (summer) to 9.5 (winter) MJ/m2 d delayed FI by between 4
and 10 d (Rawson et aI., 1984) and shorter delays were found by Marc and
Palmer (1981) over a narrower range of irradiance. Exposure to prolonged water
stress may delay FI (Marc & Palmer, 1976; cf., Yegappan et aI., 1980), while
increasing CO 2 levels to 500 Ilmolimol air may promote it (Marc & Gifford,
1974). The effect of CO 2 was limited to a short period centered about 8 d before
FI and may be linked to the cytological and physiological changes in the apex at
that time. Low N availability also may delay FI (Steer & Hocking, 1983; Hocking
& Steer, 1989) but effects of crop density appear to be limited (Villalobos et aI.,
1994). Exposure to red or far-red radiation at the end of the day determined a 12-
d difference in achievement ofFI (Connor & Sadras, 1992), a finding that may
have implications for the extrapolation of results of controlled-environment
experiments to the field.
Floral Initiation to First Anthesis
The duration ofFI to FA comprised approximately 38% of the growth cycle

of Suncross 150 over a range of constant temperatures under high irradiance
(Rawson et aI., 1984) and between 28 and 37% of the thermal duration for four
hybrids of varying maturity grown in the field (Villalobos et aI., 1994). The dif-
ferentiation of the inflorescence, up to the appearance of floret primordia in the
center of the disc, is completed in the first half of this phenophase, e.g., 17 d in
the plants grown at 28°C by Marc and Palmer (1981). During the second half, flo-
rets complete their development and attain competence to flower. The duration of
FI to FA is affected by temperature (Rawson et aI., 1984), photoperiod (Marc &
Palmer, 1981; Rawson & Hindmarsh, 1982; Rawson et aI., 1984), and cultivar
(Marc & Palmer, 1981; Rawson & Hindmarsh, 1982; Villalobos et aI., 1994),
although in some cultivars the effect of photoperiod appears to be limited (Goyne
& Hammer, 1982; Sadras & Hall, 1989).
Part of the cultivar effect on the duration ofFI to FA appears to be associ-
ated with differences in photoperiodic sensitivity. Exposure to short days hastens
development in some, but not all, cultivars (Marc & Palmer, 1981; Rawson &
Hindmarsh, 1982), and some of this effect may be due to shortening the duration
of inflorescence development, i.e., from FS 1.3 to 7 (Marc & Palmer, 1981). This
short-day response exhibited by some cultivars contrasts with long-day respons-
es found for the same cultivars during E to FI (Rawson & Hindmarsh, 1982).
These opposing responses to photoperiod during E to PI and FI to FA can give
some cultivars apparently day-neutral or even short-day responses when the dura-
tion ofE to FA is measured (Rawson & Hindmarsh, 1982). They also may con-
tribute to the ambiphotoperiodic response that has been reported for some sun-
flower cultivars (Goyne & Schneiter, 1987), although other interpretations of the
latter result are possible. Rawson et al. (1984) have suggested that shortening of
the duration between FS 1.3 and 7 under short days could be involved in reduc-
ing potential seed number.
A further source of cultivar effects on the duration of FI to FA may be
linked to the duration of E to FI. Leaf unfolding progresses at a slower rate than
leaf primordium differentiation with the result that primordia accumulate at the
apex until Fl (Yegappan et aI., 1980). Cultivars with an extended juvenile phase,
or experiencing delayed FI due to exposure to unfavorable photoperiod, are like-
ly to accumulate most. Because leaf emergence rate appears constant after Leaf 6
(Villalobos & Ritchie, 1992), duration of Fl to FA also could depend upon the
number of primordia accumulated at FI. The data of Villalobos et al. (1994) show
a positive relationship between the durations ofE to FI and Fl to FA in four cul-
tivars of increasing cycle length, a result that is consistent with the above pro-
posal.
Few attempts have been made to quantify developmental responses of sun-
flower from FI to FA (Villalobos et aI., 1996). Most early studies combined the
response within the longer phase E to FA (e.g., Keefer et aI., 1976) while others
(Goyne & Hammer, 1982; Goyne et aI., 1989, 1990) used BV to define subphas-
es of distinct environmental response. Subsequent research, discussed above,
now establishes Fl as the more significant phenostage and the better single basis
of subdivision for future work on phenological control during E to FA. An impor-
tant part of that choice lies in the different photoperiodic responses that have been
identified before and after Fl, and the possible "carryover" effects of leaf pri-
mordia accumulation at the apex. The responses considered in these quantitative
analyses do not attempt to include the full complexity of the established devel-
opmental responses, they also are variable being limited in some cases to tem-
perature (e.g., Keefer et aI., 1976; Goyne et aI., 1990), while others also include
photoperiod (e.g., Goyne et aI., 1977; Hammer et aI., 1982; Villalobos et aI.,
1996).
Despite the uncertainty surrounding the phenological control of sunflower
before and after Fl, the present, admittedly imperfect, phenological models can
be quite accurate in predicting the duration of E to FA using temperature, pho-
toperiod (where appropriate), and cu1tivar (Sadras & Hall, 1989; Goyne et aI.,
1990; Villalobos et aI., 1994a). Those relationships also have established the rel-
atively greater impact of temperature over photoperiod in the control of develop-
ment to FA, and have identified issues that need further research. In this way, they
have not only improved understanding, but also have provided powerful tools for
analysis of crop adaptation.
First to Last Anthesis
Flowering in sunflower commences at the periphery of the head and pro-

gresses inwards at up to 4 rows/d, depending upon temperature. At an average
day/night temperature of 20°C the process takes about 10 d (Rawson, 1984;
Goyne et aI., 1977) and is little affected by density or cultivar (Villalobos et aI.,
1994), although this may change if heads are very small. Each floret opens early
in the morning and its five, united anthers are rapidly extruded releasing their
pollen into the tube they form. Later in the day, the stigma extends through the
122 CONNOR & HALL
tube which at the same time withdraws as the staminal filaments age and desic-
cate. The result is that the pollen is pushed out of the tube by the emerging stig-
ma. By the following morning, the stigma is fully exserted with receptive sur-
faces completely exposed. Pollination and fertilization occur rapidly and the stig-
ma withers and recedes. Thus for each floret, the entire process is completed
within 2 d (Putt, 1940). Disruption to any of these steps is likely to result in infer-
tility and loss of yield potential.
The relative importance to infertility of damage during these steps has not
been closely studied in sunflower but it is known that, as in other crops, there is
a strong interaction with water status. Water demand increases with temperature,
windspeed, and low humidity. Low crop water status restricts cell extension and
therefore, the extrusion of anthers, dehiscence of pollen, and extension of the
stigma, as well as its receptivity. Each may contribute to infertility, which is like-
ly to be extreme under conditions of severe water shortage.
In addition, there is a direct effect of high temperature that operates in well-
irrigated crops. The phenomenon, known as heat blast, has been recorded when
air temperature exceeds approximately 40°C (e.g., Harris et aI., 1982). The criti-
cal temperature that determines this response is floret (ovary) temperature, which
differs from air temperature depending upon atmospheric conditions and crop
water supply. On sunny days, floret temperature of irrigated plants deviates
markedly from air temperature during much of the daylight period. Maximum
floret temperature is achieved 3 to 4 h before solar noon and can be as much as 8
to 9°C above air temperature at that time, and 4 to 5°C above maximum air tem-
perature that is achieved later in the afternoon (E.L. Ploschuk, personal commu-
nication). Floret temperature falls later in the day, but usually remains above air
temperature until late afternoon. Receptacle temperatures, in contrast, track air
temperatures more closely, remaining slightly (I-2°C) greater from mid- morn-
ing to sunset, presumably because of greater thermal inertia and more effective
evaporative cooling.
The magnitude of the heat blast effect depends upon the duration of high
temperature. A single day of high temperature may cause the loss of two to four
rows of florets, but with the relief from stress, normal flowering resumes. This
will have a small effect on yield, say a maximum loss of 5 to 10%. On the other
hand a prolonged period of several days of high temperature may cause not only
infertility of currently receptive florets but can damage immature florets also.
Under those conditions, infertility may continue towards the center of the head,
even following relief from stress. These effects are magnified in water-stressed
crops by higher head temperature and the direct effects of water stress. In the case
of water-short crops, it is difficult to distinguish the individual effects of high
temperature and water stress.
Last Anthesis to Physiological Maturity
The duration of LA to PM occupied about 28% of the crop cycle of

Suncross 150 under summer radiation and constant temperature (Rawson et aI.,
1984), and between 27% of the thermal time for 'Sungro 530' (long cycle) and
40% for Sunwheat-101 (short cycle) among the hybrids examined by Villalobos
et a!. (1994) under irrigation in the field. The patterns of seed growth and its com-
ponents are considered in a later section, here we concentrate on the factors deter-
mining the duration of seed filling, i.e., LA to PM.
The interval between fertilization and maximum seed weight is referred to
as the duration of seed growth. The reports of Rawson et a!. (1984), Tnipani et a!.
(1988), and Villalobos et al. (1994, 1996) show, or may be used to infer, that both
temperature and/or cultivar affect its duration, although PM always was not deter-
mined by changes in seed weight in those experiments. The duration of seed
growth also may be affected by irradiance (Rawson et a!., 1984), and it may
(Anderson et a!., 1978; Hall et a!., 1985) or may not (Whitfield et a!., 1989) be
shortened by drought. It seems likely that exposure to severe stress would short-
en seed filling if whole-plant senescence is hastened. Florets in the central por-
tion of the head reach anthesis later than those at the periphery, and if the dura-
tion of seed filling were constant across all positions, seed derived from inner flo-
rets should reach maturity later. However, this has not been clearly established,
and some results (e.g., Whitfield et a!., 1989) suggest that seed at inner positions
may fill for shorter periods than seed closer to the periphery. Effects of this nature
could reduce the position-related differences in the timing of PM with respect to
FA and LA.
Detailed experimentation based on measurements of seed or embryo dry
weight accumulation is required to establish quantitative relationships between
the duration of seed growth and temperature. It is likely that this will be aided by
separate consideration of the effects on the various subphases, viz., fertilization
to end of hull growth, fertilization to start of embryo growth, and fertilization to
start of oil accumulation (Villalobos et a!., 1996). This work should pay attention
to the thermal environment of seed. Screen temperature may substantially under-
estimate head temperature (Whitfield et a!., 1989) and changes in temperature
regime with head position and associated variations in the duration of seed
growth have been demonstrated (Ploschuk & Hall, 1995). The lengthening of the
duration of LA to PM under low irradiances found by Rawson et al. (1984) also
needs further study.
WATER RELATIONSHIPS
The Terrestrial Environment
Plants live in the interface between transiently wet soil and a relatively dry
atmosphere. To grow, they fix solar energy in C-based compounds by photosyn-
thetic reduction of CO 2 absorbed from the atmosphere, mostly by leaves. Leaves
have evolved as efficient structures for the process. They are typically planar
organs that, per unit of biomass and hence chlorophyll content, present a large
area for the interception of solar radiation and a short path length for the diffu-
sion of CO 2 from the atmosphere to the internal sites of fixation. The healthy sun-
flower plant provides a classic example of this characteristic of terrestrial plants.
The morphological adaptation of leaves to efficient photosynthesis has
important consequences for plant water relationships. An open stomatal pathway
124 CONNOR & HALL
for inward diffusion of CO 2 also is a pathway for evaporation of water from with-
in moist leaves to the dry atmosphere beyond. Loss of water from leaves (tran-
spiration) is thus an inevitable consequence of photosynthesis and causes plants
to undergo diurnal sequences of water loss and recovery. That cycling will remain
within the range of physiological tolerance provided the soil is sufficiently wet
and the root system sufficiently extensive that uptake can balance the loss from
the transpiring leaf area. As the soil dries and/or as atmospheric demand increas-
es, however, supply may fall below demand so that unless plants can adjust loss
and/or uptake, internal water status may decline sufficiently to jeopardize meta-
bolic activity or even survival. In the short term, loss may be controlled by leaf
movement or stomatal closure, and in the long term by reduced leaf expansion or
increased leaf senescence. In the long term, uptake may be increased by root
exploration and also by internal osmotic adjustment that allows plants to with-
stand lower internal water status and, at the same time, to extract more soil water.
In any event, and on either time scale, less transpiration will necessarily result in
less photosynthesis.
Photosynthesis will be discussed in the next section. Here, the focus is on
characteristics of water uptake by roots, transpiration, and the importance and
consequences of internal water status on the growth and survival of sunflower.
Collection of Water by Root Systems
The ability of root systems to absorb soil water depends upon depth (root
depth, RD, m) and intensity of exploration (root length density, RLD, cm-2).
These two parameters vary with crop development and conditions of soil mois-
ture. They also interact because roots do not penetrate dry soil. However, given
that limitation, root depth and soil texture together establish the maximum soil
water-holding capacity for a crop. Sunflower is exceptional in this regard because
its roots explore soil to greater depths, commonly extending below 2 m (Bremner
et aI., 1986; Sadras et aI., 1989), than most other annual crop species, e.g., maize,
sorghum [Sorghum hieolor (L.) Moench.], and soybean [Glycine max (L.) Merr.]
(Borg & Grimes, 1986).
Root length density defines the degree of exploration within the root zone.
The greater the density, the shorter the distance for water to move from bulk soil
to root surfaces, and hence the more effective the root system in absorbing water
from drying soil. The biomass partitioned to root growth converts to root length
via specific root length (length per unit biomass). The greater the specific root
length, the thinner the roots and the greater the root surface area per unit root
mass. Root surface area is a more important determinant of absorptive capacity
of root systems than RLD.1t is acknowledged as such in the mathematical expres-
sions that have been developed to explore the functioning of roots (e.g., Cowan,
1965) but is infrequently measured. At present, RLD is the more common basis
for quantification of the degree of root exploration.
In common with most crop species, RLD in sunflower decreases markedly
with depth, being up to 10-fold greater at 0 to 0.2 m than in deeper soil layers
(Maertens & Bosc, 1981, p. 3-11; Sadras et aI., 1989), but water availability may
change the vertical pattern of root distribution. There is little detailed information
on horizontal distribution, but Meinke et al. (l993a) found that water extraction
may continue for prolonged periods at slower rates in structured soils, possibly
due to clumped distributions of root axes. Preanthesis drought reduced mean
RLD but increased the proportion of roots in deep soil layers mainly by reducing
the amount in the surface soil of a sunflower crop (Connor & Jones, 1985) (Fig.
4-2). Interestingly, the opposite response was recorded for root distribution in the
wild sunflower H. petioiaris Nutt. (Sobrado & Turner, 1986).
Sunflower has small RLD in comparison with other crops, e.g., sorghum
and maize (Mason et aI., 1983), and shares this characteristic with legume crops
(Hamblin & Tennant, 1987). However, in both those comparisons, the species
with small RLD (sunflower and legumes) showed at least the same ability to
extract water as the mono cots indicating that smaller RLD was compensated by
more effective distribution of roots in the profile and/or by greater uptake per unit
root length. The latter could arise from lower root water potential ('l'r) or small-
er resistance to uptake across the root surface.
Sunflower's root system is thus "explorative" of large soil volumes with a
combination of thick and thin roots, small average specific root length, and small
RLD. This combination of characters enables sunflower to extract more water
than most other crops, especially from deep soil layers. This plays an important
role in productivity and in survival under low rainfall (Mason et aI., 1983;
Gimenez & Fereres, 1986; Cox & Jolliff, 1987; Hattendorf et aI., 1988; Bremner
& Preston, 1990; Meinke et aI. , 1993a). This contrasts with the "exploitative"
root systems of cereals which are characterized by predominantly fine roots, large
specific root length, and large RLD (Boot, 1990).
Some wild Helianthus spp. and several primitive genotypes are able to
develop adventitious roots at the base of stems (Rogers et aI., 1984) that are effec-
tive in water and nutrient uptake (Stevenson & Boersma, 1964). It has been pro-
posed that they contribute to the physical stability of plants and also to tolerance
to flooding . However, the production of adventitious roots is uncommon in com-
mercial cultivars and there has been no evaluation of their possible contribution
to the water or nutrient economies of crops.
1.0
0.0
0.4
]: 0.8
g
.<::
1. 2
1.6
1.8 Ra lnfed IT6) Weekly irrigated IT 1)
Fig. 4- 2. Comparison of the profiles of root length density at 80 d after sowing for sunflower irrigat-
ed weekly or rainfed. The data are the means of two cultivars, Hysun 31 and Sungold, which
responded similarly (Connor & Jones, 1985).
126 CONNOR & HALL
Plant Water Status
With the beginning of each diurnal cycle at dawn, transpiration rapidly

establishes a water potential ('V) gradient that draws water from internal tissues
and, via the root system, from the soii. The water content of crops is small so that
uptake from the soil is essential to meet transpiration loss and maintain plant 'V
within tolerable limits. Once across the endodermis of the root, where the major
hydraulic resistance occurs, direct flow of water to the canopy is relatively rapid
in the xylem, but lateral exchanges also are established from it with adjacent tis-
sues en route, depending upon their water status and resistances to transfer.
Evaporative demand falls towards dusk when transpiration may effectively cease
with the closure of stomata, but uptake continues as all tissues rehydrate towards
the equilibrium condition, the water potential of the root zone ('Vs). That condi-
tion may be achieved quickly in moist soii. As the soil dries, however, recovery
takes longer into the ensuing night period, and may not be achieved under dry
conditions. Measurements of leaf water potential ('V/) at dawn are therefore
important in studies of plant water status because they define the base level from
which plant water status can only decrease during the ensuing day.
Environmental and soil factors aside, 'Vt. and hence the photosynthetic per-
formance of the canopy, depends upon the hydraulic conductivity of the plant
(Lp). This is defined as the ratio of transpiration to ('Vr-'VD, the difference in 'V
between root surface and canopy. Measurements are difficult and seem to show
that Lp increases with transpiration (e.g., Black, 1979a,b; Turner, 1981; Koide,
1985a) but there are contrary results (Neumann et ai., 1974; Hernandez & Orioli,
1985). Even though root resistance of sunflower is small compared with sorghum
(Bremner et ai., 1986), it still contributes the major part of Lp (Boyer, 1971;
Black, 1979b; Koide, 1985a) and has been shown to vary with genotype
(Hernandez & Orioli, 1985). Taken together, these observations suggest an
important regulatory role for roots in the collection of water by sunflower and the
need for further study. New techniques of sap flow measurement (Baker & Van
Bavel, 1987) in intact plants offer much promise for studies of water flow in
herbaceous plants, including sunflower. Ham and Heilman (1990) investigated
the characteristics of such heat-balance gauges suitable for measurement of high
flow rates in sunflower stems, and Impron (1994) has successfully used them to
measure transpiration under field conditions.
Regulation of Transpiration
Two sets of responses, each appropriate to different time scales, can reduce
transpiration in the face of water shortage. Leaf movement, including wilting,
that reduces energy load, and stomatal closure that increases the resistance to
gaseous loss, are reversible responses suitable for adjustment during diurnal
cycles. In contrast, smaller leaf area, achieved by slower expansion or increased
senescence, are valuable responses to adjust transpiration over time scales of days
to weeks. Reduced expansion early in the life of the crop may be compensated
later when adequate water supply returns, but accelerated senescence is an irre-
versible loss of growth potential.
Understanding how sunflower responds to water shortage relative to other

crop plants and to explore those opportunities that exist for modification and
improvement by breeding and management is important. It is convenient to deal
with these responses in the two major phases of development, before and after
anthesis.
Pre-Anthesis
During the vegetative phase, sunflower, with its explorative root system,
relies more on restricting interception of radiation and hence evaporative demand
than it does on stomatal closure to maintain plant water status within tolerable
limits. The nature of this strategy can be seen in the extreme responsiveness of
leaf expansion to water shortage and the ability of sunflower to maintain C assim-
ilation during diurnal cycles of afternoon wilting and nighttime recovery.
The sensitivity of leaf expansion to water deficit has been demonstrated in
a number of studies. The result is that crops subjected to water deficits before
anthesis develop small leaf area index (LAI) but maintain activity per unit leaf
area (Connor & Jones, 1985; Cox & Jolliff, 1986; Guiducci, 1988; Sadras et aI.,
1991a, b). Aspects of the response of leaf expansion to water supply are present-
ed in FigA-3 in a comparison of the spatial (Fig. 4-3a) and temporal (FigA-3b)
distributions of leaf size in sunflower crops subjected to four regimes of irriga-
tion. The data show the marked differences in leaf area profiles that develop in
response to water supply (Fig. 4-3a) and also how leaves currently undergoing
expansion, but more particularly new leaves (Fig. 4-3b) are the ones to respond
following the relief from stress.
Wilting is observed most often in crops at the end of dry periods in wet-dry
cycles (Connor & Jones, 1985; Guiducci, 1988). In the experiment of Guiducci
(1988), the fraction of radiation intercepted during the season ranged from 83 to
33% in crops that were irrigated weekly, biweekly, or were rainfed. The effect of
wilting was evident in a "saw-toothed" pattern of radiation interception, particu-
larly of crops irrigated biweekly.
Leaf conductance (g/) is dominated by the stomatal component (gs) because
the parallel conductance through the cuticle is small. Changes in gl can therefore
be taken to reflect those in gs' It has been shown that sunflower stomata are less
sensitive to water deficit than in other species (Sionit & Kramer, 1976; Connor &
Cawood, 1978; Goudriaan & van Laar, 1978; Rawson, 1979; Sobrado & Turner,
1983a, b; Connor & Jones, 1985). Before anthesis, no changes in gl were found
in young leaves of stressed crops (\jIj, approximately -1.6 MPa) in comparison
with irrigated controls (\jIj, from -1.0 to -1.2 MPa). During this period, water-
short crops seem able to maintain gl at levels similar to those of well-watered
crops by other adjustments (Sadras et aI., 1991 b, 1993c). Even in crops that are
visibly wilted, stomata do not close completely and photosynthesis continues
(Connor et aI., 1985b).
There is debate about the physiological responses that control leaf expan-
sion and gs in plants, and sunflower has figured strongly in that research. The
alternatives, not necessarily exclusive, are a hydraulic feedback response from \jI1
and a feed-forward signal from the roots that may sense water depletion in
128 CONNOR & HALL
advance of low"" (Davies & Zhang, 1991). A number of experiments with sun-
flower, mostly performed on potted plants grown under controlled conditions,
suggest abscisic acid (ABA) as a likely candidate for a "root-signal" compound
(Ehret & Boyer, 1979; Robertson et aI., 1985; Hubick & Reid, 1988; Neales et
300 T4 anthesis
250 first
Days after sowi n9
Fig. 4--3. (a) Leaf area profiles of sunflower Sungold under four regimes of irrigation. Tl and T6 are
the weekly irrigated and rainfed controls. In T4 and T5, weekly irrigation was commenced at late
budding (50 d after sowing, DAS) and anthesis (DAS 70), respectively. (b) Patterns of expansion
of individual leaves, numbered from the base, of sunflower Sungold under the various irrigation
treatments described for Fig. 4--3a (after Connor & Jones, 1985).
aI., 1989; Zhang & Davies, 1990). A recent study (Sadras et aI., 1993b) has
attempted to identify the mechanisms of response of sunflower crops in the field.
It used a series of experiments including split-root treatments in which root sys-
tems of individual plants were subjected simultaneously to wet and dry soil con-
ditions. Leaf expansion was closely correlated with 'VI across all treatments inde-
pendent of the soil moisture and environmental conditions under which it devel-
oped. The authors could not, however, discount the importance of possible inter-
play between hydraulic and nonhydraulic effects on gs which have been observed
in small plants grown in pots (Gollan et aI., 1992; Schurr et aI., 1992). These lat-
ter effects appear quite complex, with both ABA and stomatal sensitivity to ABA,
among the possible factors involved.
Post-Anthesis
Leaf expansion is complete by anthesis so that subsequent changes in LAI
result only from leaf senescence. Increased leaf senescence is less effective than
reduced leaf expansion in regulating LAI and transpiration for two reasons. First,
senescence is less sensitive to soil water deficit than is expansion (Sadras et aI.,
1991 b). Second, senescence reduces leaf area from the base of the canopy where
leaves contribute relatively little to total transpiration because of lower irradi-
ance, small individual area and small gI (English et aI., 1979; Rawson et aI.,
1980).
Stomatal closure appears to playa greater role in the control of plant water
status after anthesis. Connor and Jones (1985) showed that in crops stressed after
anthesis, gI fell to 50% of the irrigated controls. A similar response also was
reported by Cox and Jolliff (1987). The impact of wilting on transpiration of
crops stressed after anthesis seems to be similar to that discussed previously for
preanthesis stress (Guiducci, 1988).
Variation with genotype in the effect of water deficit on gI (Hernandez &
Orioli, 1985; Gimenez & Fereres, 1986) and Lp (Blanchet et aI., 1978, p. 12-22)
makes it difficult to generalize about the effect of changes in gI on transpiration
in sunflower. The response of stomata to osmotic adjustment (Turner et aI., 1978;
Conroy et aI., 1988) together with the large intraspecific variability in osmotic
adjustment in this species (Chimenti & Hall, 1993) suggest that intraspecific vari-
ability in gI is greater than was previously thought (e.g., Turner et aI., 1978).
In summary, crops stressed before anthesis regulate transpiration predomi-
nantly by reduced leaf expansion whereas after anthesis it is difficult to identify
a dominant factor. Rather, there is an interplay between reduced interception by
hastened leaf senescence, wilting, and reduced gs' More work at the crop level is
necessary for an accurate determination of the relative effect of each of these fac-
tors on crop transpiration under various timings and intensities of water deficit. It
is worthwhile concluding this section by emphasizing that the lack of response of
gs to water shortage in young sunflower plants does not support the argument that
the species exerts little control over water use (e.g., Rawson & Munns, 1984). To
accept that would be to ignore the effective regulation of transpiration achieved
by the marked sensitivity of leaf expansion to soil water deficit.
130 CONNOR & HALL
Drought Tolerance
Sunflower is considered well adapted to drought, although systematic

analyses of the physiological basis of it, and/or purposeful attempts to breed for
greater drought tolerance are limited. Important issues here are that drought tol-
erance is not a simple attribute, that patterns of water availability (and hence,
likelihood, and nature of droughts) vary widely between agroecosystems, and that
individual combinations of physiological attributes effective under one set of cir-
cumstances may not be under others. Management decisions, through selection of
cultivars and sowing dates, are important in this context. Thus, shifting the crop-
ping season to less evaporative periods of the year may reduce exposure to
drought (e.g., Gimeno et ai., 1989), while use of early maturing cultivars may
increase the fraction of water used after anthesis leading to greater harvest index
(HI) and yield (Sadras & Connor, 1991) provided water extraction from the pro-
file is complete (Fereres et ai., 1986) and crop population density is adjusted to
ensure adequate exploitation of available resources by early maturing cultivars
(Villalobos et ai., 1994).
Plant breeders have made progress in matching cultivar characteristics to
environment and this has probably included some unconscious selection for
drought tolerance. The question that remains is whether a structured approach,
which includes the identification, testing, and breeding for particular attributes,
could increase the rate of progress. However, the challenge is great in regions
with marked interyear variability in rainfall-a common characteristic of
drought-prone areas. Useful reviews of these issues can be found in Ludlow and
Muchow (1990) and Passioura (1986). Here we briefly review work on sun-
flower.
The most extensive comparative analysis of drought tolerance of sunflower
cultivars is that of Fereres et ai. (1986) and Gimenez and Fereres (1986). They
compared a range of genotypes over several years, growing crops under both irri-
gated and rainfed conditions that produced terminal droughts. Importantly, they
found no association between yield potential and susceptibility to drought with
important implications for the development of high-yielding, drought-tolerant
cultivars (see also Baldini et ai., 1992a; Barron, 1992). They also showed that
yield under drought was closely associated with HI, and found intraspecific vari-
ability for rooting depth, which in tum was linked to cultivar maturity type.
Further, they identified variability for both sensitivity of leaf conductance to leaf
water potential and for osmotic adjustment. In extrapolating these results to vari-
able, low-rainfall environments it is important to realize the importance of timing
and probability of water shortage. In any environment, high-yield potential is
associated with long growing season which itself increases the risk of low HI
under drought (Turner & Rawson, 1982).
There is good correlative evidence that osmotic adjustment is a trait which
may confer drought tolerance in crops such as wheat (Triticum aestivum L.
emend. Theli.) and sorghum (Morgan et ai., 1986; Santamaria et ai., 1990).
Chimenti and Hall (1993) have found significant intraspecific variability for this
trait in sunflower and its narrow-sense heritability is substantial (Chimenti et ai.,
1992). Interestingly, genotypes with a high capacity for osmotic adjustment are
those that are more restrictive of leaf expansion under drought (Chimenti & Hall,
1994). This response can contribute to yield maintenance depending on whether
continued growth or survival has the greater impact.
Wild sunflower species are a possible source of traits which confer drought
tolerance and some effort has gone into examining the comparative water rela-
tionships of wild and cultivated sunflower species (e.g., Sobrado & Turner,
1983a,b; Morizet et aI., 1984; Serieys, 1991; Baldini et aI., 1992a,b). Several
putative traits have been identified but understanding of causal linkages with
yield maintenance is, at best, weak. While it would be inappropriate to ignore the
consistent link identified between drought tolerance and osmotic adjustment in
other species because causality has not been clearly established, it also is clear
that a good understanding of the mode of action of individual traits will increase
the chances of their successful use in breeding programs. This is particularly true
where the indicator used is itself the end-product of complex interactions. Thus,
maintenance of leaf water content under drought (e.g., Baldini et aI., 1992b) or
low canopy temperature may reflect several different interactions between root-
ing depth, stomatal sensitivity, and leaf area.
In summary, the evidence suggests that it should be possible to breed for
tolerance to at least some categories of drought in sunflower without loss of yield
potential because there are firm indications of intraspecific variability in traits
which can confer tolerance, and some information on heritability. The likelihood
of success will be increased if work is based on a good understanding of the
causal linkages between the possession of a trait and the physiology of yield loss
minimization under stress (Ludlow & Muchow, 1990).
MINERAL NUTRITION
Sunflower shares with most higher plants the essential requirement for six
macronutrients (N, P, K, Ca, Mg, and S) and seven micronutrients (Fe, Mn, Zn,
Cu, B, Cl, and Mo) (Blarney et aI., 1987). Uptake depends upon root exploration,
soil water content, and the available, rather than total, nutrient content in the soil.
Nutrient elements combine in precise stoichiometric relationships in metabolic
reactions but chemical composition of biomass is not fixed for two reasons. First,
some functions can be achieved by alternative nutrients, e.g., osmotic balances of
cells that are critical to turgor, can be achieved by a variety of ionic combinations.
Second, plant functions provide for storage, mobilization, and reuse of nutrients.
This section concentrates on issues that distinguish sunflower nutritional
physiology from other plants. Management of crop nutrition with fertilizers is
discussed in Blarney et al. (1997, Chapter 12).
Micronutrients
Micronutrients are so called because they are required in small amounts and
hence are infrequently deficient in crop production. When recognized, deficien-
cies are readily and inexpensively corrected with fertilizer. Availability of indi-
vidual nutrients responds differently to soil pH. Increasing pH can induce defi-
132 CONNOR & HALL
ciency of Fe, Mn, Cu, and Zn or toxicity of B and Mo. Sunflower was able to
extract more Fe from a deficient, calcareous soil than sorghum because of a
greater acidifying effect on soil in the immediate vicinity of roots (Mathers et aI.,
1980).
Sunflower has a large requirement for B compared with other plants and
has been used as a test species for B availability in soils (Schuster & Stephenson,
1940). Macroscopic symptoms of B deficiency generally appear around flower-
ing and include brittle stems and peduncles as well as malformed leaves and
inflorescences. Optimum leaf concentrations have been reported in the range 30
to 60 mg/kg. In comparison, Gonzalez Fernandez et ai. (1985) reported stem
breakage and head loss due to B deficiency in plants with concentration in the
upper expanded leaves of 10 to 13 mg/kg. In less severe cases, malformed heads
produced many nonviable seed, often concentrated in patches. These symptoms
are consistent with cytological effects that can be observed in deficient plants ear-
lier in crop development in the form of impeded cell division and depleted DNA
in the cells of the disc-floret, generating zone of the receptacle during floret ini-
tiation (Palmer et aI., 1988).
Blarney et ai. (1979) reported intraspecific variability in response to B sup-
ply, part of which may be related to varying ability to extract B from deficient
soils (Blarney & Chapman, 1982). Mycorrhizal associations (see subsequent sec-
tion) may enhance B uptake from deficient soils (Jodice et aI., 1981) and thus
contribute to differential responses between cultivars.
Macronutrients
The uptake and distribution of the three major macronutrients, N, P, and K,

have been studied in sunflower by Gachon (1972) and Mathers and Stewart
(1982). Figure 4--4, adapted from Gachon (1972), serves to emphasize the fol-
lowing important issues in the physiology of plant nutrition: (i) considerable
quantities of macronutrients are required to support crop growth, (ii) the greater
proportion is absorbed in the first half of the growing season, (iii) the second half
of the season is characterized by redistribution of nutrients among component
organs, (iv) each nutrient has an individual pattern of uptake and redistribution,
and (v) different proportions of each nutrient are removed by harvest of seed.
Potassium
Potassium has its major role in the osmotic balance of cells and hence in
the maintenance of turgor and cell extension. In this way, it has a direct effect on
the growth of all plant organs, particularly stems, leaves and roots.
Photosynthesis and transpiration, also respond indirectly to K availability
because it is the principal cation involved in the control of guard cell turgor and
hence the opening and closing of stomata (Ehret & Boyer, 1979). In addition, at
the metabolic level, K functions as a cofactor of many enzyme-catalyzed reac-
tions (Taiz & Zeiger, 1991).
The distribution of K within the plant reflects its association with active
cellular expansion. During growth, large amounts of K pass in sequence from
petioles to leaf laminae, stems, and finally receptacles (Gachon, 1972). In con-
trast, [K] of seed is small, in the range 5 to 10 g/kg (Mitreva, 1989; Robinson,
1973; Seiler, 1986a), amounting to 5 to 10 kg/(t seed). Expressed another way,
only around 5% of the considerable quantity (140 kg/ha) absorbed by crops yield-
100
I
I
Nitrogen I
I
80 lamina I
I
stem I
receptacle I
I
petiole I
60 seed I
I
I
I
I
'"
I
.<:
OJ 40
"'"
20
o.
20 40 60 80 100 120140
Days after sowing
160
140 Potassium
120 '~-' ...

100
'"
.<:
Cl
80
""'- 60
40
20
//./
0
0 20 40 60 80 100 120 140
Days after sowing
15
I
I
Phosphorus "
I
I
I
I
I
10 I
I
I
I
I
I
I
I
I
I
I
-"
I
//z::':~;:~::
20 40 60 80 100 120 140
Days after sowing
Fig. 4-4. The concentrations and distribution ofN, P, and K among aboveground organs of sunflower
(after Gachon, 1972).
134 CONNOR & HALL
ing 1 tlha (Sfredo et aI., 1983) is removed in the seed. The major part ofK accu-
mulated by the crop is returned to the soil in the vegetative parts at harvest.
Nitrogen
Nitrogen is the macronutrient required in the greatest quantity by plants. In
sunflower, concentrations in stems, leaflaminae, and seed may rise to around 20,
30, and 40 g/kg, respectively. Nitrogen is an active component of many metabol-
ic and storage compounds but has a major association with photosynthetic
enzymes that occur in large quantities in leaves. In C3 species generally, three-
quarters of the leaf N content is associated with photosynthesis (Evans, 1989),
and in sunflower, the single photosynthesis enzyme, rubisco, accounts for around
50% of soluble leaf protein (Gimenez et aI., 1992).
Nitrogen may exist in the soil or be applied as fertilizer in various forms but
all are rapidly transformed to nitrate under field conditions where it exists in the
soil solution. Sunflower roots contain only small amounts of the enzyme nitrate
reductase so absorbed nitrate is mostly translocated directly to the leaves via the
xylem (Kaiser & Lewis, 1980; Hocking & Steer, 1983b). There, in association
with photosynthesis, it is reduced to amino acids, some of which are translocated
in the phloem to sites of active growth around the plant (Merrien et aI., 1988), but
large amounts of N remain in the photosynthetic system of expanding leaves.
Later, as leaves senesce, N is mobilized from rubisco and translocated to expand-
ing leaves and, as the crop matures, to seed. Root systems remain capable of N
uptake during seed filling so that fertilizer or foliar sprays during that period slow
the rate of leaf senescence and may increase yield (Go swami & Srivastava,
1988). The response of photosynthesis to N is considered in the next section.
Shortage of N affects the development and growth of both source (leaves)
and sinks (florets and seed). Nitrogen deficiency during the early vegetative peri-
od may reduce the number of leaves, but it will certainly restrict their expansion
(Steer & Hocking, 1983; Hocking & Steer, 1989; Connor et aI., 1993, Gimenez
et aI., 1994). This is shown in the comparison ofleaf area profiles of crops grown
at two contrasting levels ofN supply in Fig. 4-5 (Gimenez et aI., 1994). The
result is reduced interception, which together with a likely smaller photosynthet-
ic rate, greatly reduces crop growth rate.
Mobilization of N within the plant concentrates about 60% in the seed
(Gachon, 1972). Nitrogen content of seed is usually in the range 25 to 35 g/kg so
that a yield of 2 tlha removes up to 70 kg N/ha. Crop management must general-
ly seek to replace that to maintain the yield of subsequent crops. The relative
importance of mobilization and uptake as sources of seed N vary during seed fill-
ing, among other factors, with the level of soil N. For instance, mobilization and
uptake were 43 and 57%, respectively, in plants with high-N supply (Hocking &
Steer, 1982) and 75 and 25% in plants with N deficit (Vrebalov, 1974). Leaves
are the major source of mobilized N so the photosynthetic system serves as the
source of both carbohydrates and N compounds for the developing seed. Steer et
al. (l985a,b) compared the response of seven genotypes to N supply and con-
cluded that genotypes with the greatest seed yield combined effective redistribu-
tion from vegetative parts with continued uptake from the soil after anthesis.
15----------------------~
(a)
10
Q;
-"
E
:::I
c:
1ii
Q)
...J
5
DAS 33
0
50 100 150
Leaf area (cm 2)

30
20
Q;
-"
E
:::I
c:
Oi
Q)
...J 10
o+-~--~~--~----~--~
o 200 400 600 800
Leaf area (cm 2 )
Fig. 4-5. Profiles of leaf area of well-watered sunflower crops (Sungro 385) at Cordoba, Spain, on
two occasions: (a), 33 and (b), 63 d after sowing (DAS), in response to N. Crops were sown at two
densities (2.9 m~i! closed symbols and 5.7 m'" open symbols), each was grown with (circles) and
without (squares) added N (25 g/m2) (Gimenez et aI., 1994).
Phosphorus
Phosphorus is required in smaller amounts than either K or N. In adenosine
triphosphate (CIOH1SNsOIOP, ATP), P plays the central role in energy transfers for
chemical synthesis and so is an essential component in all metabolically active
cells. Phosphorus also is a component of the phospholipids in cell membranes
and of nucleotides. Although uptake of P continues during seed filling, if it is
available in the soil (Vrebalov, 1974; Hocking & Steer, 1983a), lack of soil mois-
ture may severely limit uptake by rainfed sunflower during this period. As with
N, there is extensive mobilization ofP within the plant (Hocking & Steer, 1983a),
and this can be an important source of P for seed development under conditions
of restricted P uptake. Estimates of the contribution ofP mobilized from the stem
and leaves to mature seed range from about 30 (Hocking & Steer, 1983a) to over
60% (Vrebalov, 1974). There is considerable accumulation of P in the lipo-pro-
136 CONNOR & HALL
teins and phytate of seed. Hocking and Steer (1983a) estimated that 80% oftotal
plant P accumulates in seed and field experiments (Gachon, 1972; Sfredo et aI.,
1983) show that up to 75% of crop P is removed in seed.
Unlike N, P is immobile in soil so that uptake depends upon thorough
exploration by the root system. In sunflower, as in a range of other species, that
is aided by symbiotic association with vesicular arbuscular mycorrhizae (VAM),
in sunflower formed with the fungal genus Glomus (family Endogenacae) (Hall,
1984). The fungal hyphae provide considerable "extension" to the root system
allowing infected plants to take up immobile nutrients otherwise inaccessible,
physically and probably also chemically. The degree of mycorrhizal infection in
sunflower appears to be inversely related with soil P level, with infection affect-
ing leaf expansion and biomass partitioning in seedling plants (Koide, 1985b). It
is clearly important to extend these studies to encompass yield responses of
crops.
CARBON ASSIMILATION
Biomass accumulates in C compounds achieved as the balance between

photosynthesis and respiration. Photosynthesis, mostly by leaves but also by
stems and capitula, is restricted to daytime when it varies sinusoidally with solar
irradiance. In contrast, all organs respire continuously.
Chemically, respiration is the breakdown of glucose with the release of
simpler C skeletons, energy, and CO 2, Those skeletons are used in a wide variety
of chemical reactions, supported by the energy released. It is useful, however, to
identity two functional groups of processes. The first, known as maintenance res-
piration, sustains the structural and metabolic integrity of existing membranes,
cells, and tissues. The second, growth respiration, sustains the synthetic process-
es of the formation of new biomass.
As a set of biological reactions, respiration is sensitive to temperature with
an approximate doubling for each 10DC rise in temperature. Thus maintenance
respiration (MR) can be expressed as MR = W.RM.QIO.exp[(T - 20)/10], where W
is organ size, RM is the rate of maintenance respiration per unit size at 20 DC, and
T is temperature (DC) (Loomis & Connor, 1992). The coefficient, QlO, is the
increase in rate for each IODC rise in temperature. As a first approximation, QIO
= 2 (a doubling) is appropriate, but empirical estimates for individual organs
under a range of conditions varied between 1.4 and 2.2 (Hall et aI., 1990a). The
mean value for heads was 1.71 and compares with a value of 1.84 reported earli-
er by Horie (1977).
The rate of growth respiration also depends upon temperature but equally
it can be related to product type. Carbohydrates, proteins, lipids, and lignin
require distinct amounts of energy per unit biomass synthesised. Calculations can
be made either by consideration of each chemical step in synthesis or by elemen-
tal analysis of the products (Loomis & Connor, 1992). The result is a fixed
growth respiration per unit of product type, ranging from 0.12 g C0 2/g carbohy-
drate to 1.61 g C0 2/g lipid. Alternatively, growth respiration can be expressed as
a production value (PV), i.e., gram product/gram glucose required (0.83 for car-
bohydrate and 0.37 for lipid). Growth proceeds faster at higher temperatures but
the energy cost of growth remains the same.
This section deals with development of leaf area, the interception of radia-
tion, and the photosynthesis and respiration responses of component organs and
complete canopies to solar irradiance, temperature, water, and N supplies.
Leaf and Canopy Area
The canopy area of individual plants and of crops is the product of green
leaf number and individual leaf area, which represent the outcome of the separate
processes of leaf primordium initiation, leaf appearance, expansion, and senes-
cence.
The embryo contains two leaf primordia (von Guttenberg et aI., 1955) and
further primordia are differentiated at rates which approximate 0.069 primor-
dia/°Cd (Tb = 4°C) between E and FI in nonstressed plants (Sadras & Villalobos,
1993). Exposure to severe water or N stress may reduce the number of primordia
differentiated during this phase (Marc & Palmer, 1976, 1978; Steer & Hocking,
1983) and the maximum number of primordia for individual cultivars may vary
in response to photoperiod and temperature effects on the duration to FI (Rawson
& Hindmarsh, 1982; Goyne & Hammer, 1982; Goyne & Schneiter, 1988). Effects
of radiation also have been reported (Rawson & Dunstone, 1986). However, in
most circumstances, genotype, not environment, is the major cause of variation
in total leaf number (e.g., Rawson & Hindmarsh, 1982; Goyne & Schneiter, 1988;
Sadras & Villalobos, 1993).
Leaf appearance proceeds more slowly than primordium differentiation
(Yegappan et aI., 1980) at a rate strongly dependent on temperature, but with less-
er effects of daylength and cultivar (Rawson & Hindmarsh, 1982; Villalobos &
Ritchie, 1992). The first leaves (Positions 1-6) appear more slowly than the
remainder, with values of 38.7 and 23°Cd (Tb = 4°C) per phyllochron, respec-
tively (Villalobos & Ritchie, 1992). Leaf appearance rate is reduced at tempera-
ture above an optimum around 27°C (Villalobos & Ritchie, 1992) and by heavy
shading (Rawson & Hindmarsh, 1983).
Final leaf size is strongly influenced by nodal position, temperature, and N,
water, and radiation stresses (as reviewed below). It also is sensitive to root zone
aeration (Kriedemann et aI., 1983) and salinity (Rawson & Munns, 1984).
Cultivar effects on mean leaf size have been reported (Rawson & Hindmarsh,
1982) but the main effect of cultivar seems to be associated with total leaf num-
ber and associated effects on the pattern of leaf size with node (Fig. 4-3a). Crop
population density effects on leaf area per plant are important (Sadras & Hall,
1988; Sadras et aI., 1989), and presumably include effects on both expansion and
senescence. Although these effects have not been studied in detail, final leaf size
at the uppermost three-quarters of all nodes is reduced once crop population den-
sity exceeds some threshold value (M.C. Rousseaux, personal communication).
Dale (1988) has recently reviewed aspects ofleaf expansion and its control.
Final leaf size may be regarded as the product of the rate and duration of expan-
sion. Of the two components, duration is relatively insensitive to node position,
leaf water and N status, but relatively sensitive to temperature and radiation. In
138 CONNOR & HALL
contrast, rate of expansion is much affected by leaf water and N status and node
position, as well as by radiation and temperature (e.g., Rawson & Hindmarsh,
1982; Rawson et aI., 1984; Rawson & Dunstone, 1986; Steer & Hocking, 1983;
Sobrado & Rawson, 1984; Connor & Jones, 1985; Connor et aI., 1993; Trapani
et aI., 1993). The interactions between the responses of these two components of
leaf size contribute greatly to the complexity of canopy responses to cultivar and
environment.
Temperature and radiation effects on duration of leaf expansion, which may
contribute to canopy response to sowing date, have been studied in detail
(Rawson & Hindmarsh, 1982; Rawson & Hindmarsh, 1983; Rawson et aI., 1984;
Rawson & Dunstone, 1986). Across nodes, under high levels of radiation (25.5
MJ/m 2 d) and between 15 and 27 DC, duration was close to 360 DCd (Tb = ODC). For
each MegaJoule/ square meter per day reduction from that level, duration
increased by almost 5DCd (Rawson & Dunstone, 1986). These authors also found
some indications of cultivar effects on duration of leaf expansion.
Leaf expansion is partly attributable to cell division, with palisade cell
numbers increasing 10-fold between leaf appearance and achievement of approx-
imately 30% of final leaf size. Cell expansion then continues until the completion
of leaf growth (Yegappan et aI., 1980). Leaf area, at least for the leaves below
maximum size, is positively correlated with cell number (the dominant compo-
nent) and negatively correlated with cell size (Yegappan et aI., 1982).
Both temperature and radiation have substantial effects on leaf expansion
rate. Average expansion rate for the largest leaves almost doubled between 9.5
and 25.5 MJ/ m2 d, and increased by almost 20% between 15 to 21 DC under high
radiation, although it decreased at higher temperature (Rawson & Dunstone,
1986). The important consequence of the responses of duration and rate of expan-
sion to temperature is that largest plant leaf areas are generated under cool con-
ditions (Rawson et aI., 1984; Rawson & Dunstone, 1986). The effects of radia-
tion may be linked to availability of photoassimilate for expansion, as suggested
by growth responses to shading for longer than 12 h (Takami et aI., 1982).
However, leaf size is not linearly dependent on radiation, and adaptation to low
irradiance may involve changes in specific leaf area as well as cell number and
size (Dengler, 1980; Rawson & Hindmarsh, 1983).
The effects of water stress on leaf expansion are mediated by changes in
both cell number and cell size. The latter effect predominates in leaves unfolding
early during stress episodes and the former in later stages of exposure to stress,
consistent with the partial temporal separation of the processes of cell division
and expansion (Yegappan et aI., 1982). Much work has been done on the effects
of water stress and subsequent recovery of leaf expansion. Expansion rates, rela-
tive to well-watered controls, varied little among the four cultivars examined by
Takami et al. (1981), and could largely be explained by variations in predawn tur-
gor, with expansion measured over several days increasing sharply at values
greater than 0.2 MPa. Diurnal patterns were more complex, and relationships
between expansion and turgor only held for daytime measurements (Takami et
aI., 1982). Work by Matthews et al. (1984) showed that acclimated leaves (i.e.,
leaves exposed to water stress during expansion) grew more slowly than unaccli-
mated leaves during recovery from stress, and these effects could be explained by
changes in yield threshold and tissue extensibility, with little change in hydraulic
conductance. However, responses to recovery on a whole plant basis are more
complex (Takami et aI., 1981; Rawson & Turner, 1982a,b; Connor & Jones,
1985). Leaves at some nodes grow less than their equivalent on well-watered
controls after recovery, others grow as much, and a few may grow more. Both
leaf age and position can affect these responses, with unemerged leaves furthest
from the capitulum having the greatest probability of expanding to normal or
greater than normal size on relief of stress (Rawson & Turner, 1982b). These
complex responses can substantially modify crop leaf area profiles. As vegetative
crops enter periods of stress, the currently expanding leaves are restricted. If the
stress is short-lived, they may be able to respond to some extent to relief from
stress, otherwise response is seen in a younger cohort of leaves still in their early
phases of expansion. If stress is not relieved, all leaves are much reduced in size
(Fig.4-3b).
Nitrogen availability has a profound effect on leaf size (Fig. 4-5), largely
through its effects on the rate of leaf expansion (e.g., Steer & Hocking, 1983;
Connor et aI., 1993; Trapani et aI., 1993, Gimenez et aI., 1994). However, the
physiological bases of these effects are poorly understood. Present information
suggests that the threshold and saturation levels of specific leafN for leaf expan-
sion are greater than those for photosynthesis (Trapani et aI., 1993). In contrast to
the acclimation to water stress, changes in hydraulic conductivity rather than
yield threshold or tissue extensibility underlie the responses of leaf expansion to
N (Radin & Boyer, 1982).
Of all processes contributing to canopy size, leaf senescence has received
the least attention. Associations have been demonstrated between senescence and
elapsed thermal time from both emergence and anthesis (Sadras & Hall, 1988;
Chapman et aI., 1993b), and these relationships are probably reflections of the
progress of seed filling and the consequent loss of N from leaves (Sadras et aI.,
1993a). Leaf senescence may commence before anthesis, particularly in crops of
large LAI, and there has been some research on the linkage between canopy N
profiles and canopy photosynthetic capacity (Sadras et aI., 1993a; Gimenez et aI.,
1994; Connor et aI., 1995). Rousseaux et aI. (1996) have shown that leaflongevi-
ty following maximum size may be reduced from 50 to 9 d by heavy shading, and
that there is a linear relationship between longevity and mean daily radiation dur-
ing the shading period. There is good evidence that water (e.g., Connor & Jones,
1985; Whitfield et aI., 1989) and N stresses (Sadras et aI., 1993a) may affect leaf
senescence in sunflower but little is understood of these important effects.
Interception of Radiation
Irradiance declines exponentially with leaf area within crop canopies. This
can be measured either as a rapidly decreasing average irradiance or a similar
trend in the proportion of sunlit area. Analysis of the former requires simple and
the latter more complex mathematical treatments. The discussion here will use
the simple exponential extinction profile model to discuss changes in interception
in response to crop and environmental factors. In this model, average irradiance
(I) below LAI = L of a canopy receiving an ambient irradiance (10) is given by the
140 CONNOR & HALL
expression 1= Io.exp(-kL) (FigA-6). The extinction coefficient, k, describes the

geometry of leaf display. For randomly distributed leaves, k depends mostly on
leaf angle. Because sunflower is sown in rows, this relationship may not provide
an accurate description of interception when LAI is small and horizontal distrib-
ution of leaves deviates substantially from random.
Heliotropism is a notable feature of sunflower crops that complicates the
study of interception because leaves and capitula track the sun diurnally up to
anthesis. The result is greater interception and enhanced photosynthesis. The pro-
portion of leaves with heliotropic ability that is sunlit decreases with ontogeny so
the advantage of heliotropism to crop photosynthesis is largely restricted to
young stands in which it may reach 20% (Shell & Lang, 1976).
Diurnal and seasonal patterns of k have been described for sunflower crops.
Diurnally, k decreases until solar elevation exceeds the leaf angle that in sun-
flower canopies is around 30 0 (Horie & Udagawa, 1971). When that geometry is
established, only the upper leaf surfaces are sunlit and k remains constant until
solar elevation again falls below leaf angle at the end of the day. Empirical deter-
minations show daily patterns that are consistent with this model (Sadras et ai.,
1991 b). In this context, solar tracking ensures that more upper leaf surfaces are
illuminated at low solar angles so one effect is to maintain a constant k for longer
during the day. It also is likely that the response reduces overlap as individual
leaves follow the sun. That would reduce the randomness ofleaf distribution that
characterizes static leaf canopies with a consequent tendency to increase k.
Interception of radiation also depends on crop water status. In particular
wilting, which is common diurnal behavior in water-stressed crops, leads to more
vertical leaf display and hence smaller k (Guiducci, 1992a, b). The effect of these
structural changes on canopy photosynthesis is discussed in a later section.
1.0-.----------------,
0.6
0.6
0.4
o Artlung E353
• Alhama Extra
0.2 • Florasol
o RP-4106
0.0 +--__ -r-~-r_~-__._-~___l
o 3 4
Leaf area index
Fig. 4-6. Fraction of incident photosynthetically active radiation intercepted at noon (Qnoon) as a func-
tion of leaf area index (LAI) for four cultivars of sunflower. The fitted line (,-2 = 0.98) follows the
equation Qnoon = I - exp(-0.86 LAI) (Orgaz et aI., 1992).
Seasonally, k decreases with ontogeny until about anthesis and then

increases towards the end of the cycle (Sadras et al., 1991 b). That results from
increasing overlap between leaves, within and between plants as LAI increases,
and possible changes mediated by the decreasing proportion of sun-tracking
leaves (Lang & 8egg, 1979).
Given these complications, small differences in k do exist between culti-
vars. For example, Sadras et al. (1991b) established that crops of semidwarf
hybrids had smaller k than standard-height cultivars early in the season, a differ-
ence they attributed to greater overlap between leaves concentrated on shorter
stems. In contrast, Orgaz et al. (1992) established a common mean noontime k =
0.86 (Fig. 4-6) for four cultivars. Large differences in k in sunflower could only
be achieved by considerable change in leaf angle, probably beyond the range that
exists in currently available germplasm.
Photosynthesis and Respiration
Individual Leaves
Sunflower leaves, as for other C 3 species, have an asymptotic relationship
between net photosynthesis (P n) and incident irradiance (I). The response can be
characterized in physiological terms (Goudriaan & van Keulen, 1979) as
Pn = (Pmax + R){ 1 - exp[-eI/(Pmax + R)]} - R [1]
where P max is the maximum rate of photosynthesis at saturating irradiance (/), £

is the apparent quantum efficiency (the slope, dPnldI, at I = 0), and R is the res-
piration rate in the dark. Changes in temperature affect the relationship mostly
through respiration which, as previously explained, responds exponentially to
temperature. One way to account for this in comparisons of Pn is to adjust R to a
common temperature within the broad optimum for photosynthesis, say 25°C
(e.g., Connor et al., 1993). The compensation point, which derives from the equa-
tion, is the irradiance at which P n = O.
The temperature sensitivity of sunflower photosynthesis has been the sub-
ject of several studies (Horie, 1977; Warren Wilson, 1966; Paul et al., 1990)
which define the broad optimum between approximately 17 and 32°C.
Photosynthetic activity falls sharply at lower temperature, although positive rates
are still found at 5°C (Paul et al., 1990). Prior exposure to low temperature also
can contribute to lower leaf photosynthetic rate (Paul et al., 1990), probably due
to a negative feedback resulting from accumulation of assimilate in leaves (Paul
et al., 1991).
The photosynthetic characteristics of sunflower leaves are well established
(Fig. 4-7). The maximum photosynthetic rate is large in comparison with other
C3 species and is achieved at higher irradiance. In ambient air {[C0 2] approxi-
mately 330 /lmol/(mol air)} and with temperature in the range 25 to 35°C, Pmax
of 40 /lmol COi m 2 s is achieved at irradiance exceeding about 1500 /lmol/m2 s
photosynthetically active radiation (PAR) (English et al., 1979; Connor et al.,
1993), twice the average value for C 3 species achieved around 950 /lmol
142 CONNOR & HALL
0.10
(a) a Nl
·•
>-
A N2
0
c: 0.08 N3
·0'" 0 N4
;;:::
a; D. • N5
• 0
0.06
° •
E
:>
cItS A
Dj>A",:?:
-"Ii ", •• 4> 0°<1"0
• -",:'Ii
:>
CT 0.04 • lid' A';O •
•
_• • • 0
C A •
~
ItS
a.
a. 0.02 mean 0.050 (SE 0.001)
<:
3
0.00
0 2 3
Specific leaf nitrogen (g N m-2)
50
(b)
40
•
rn
·iii
'c"
£: 30
>- U,
rn
N
B
0
.J:: E 20
a.
"0
E
:>
E
§ 2: 10
)(
ItS
::;;
·10
0 2 3
Specific leaf nitrogen (g N m- 2)
Fig. 4-7. Relationships between parame- (e)

ters of the leaf net photosynthesis ~.
response in sunflower and specific leaf 4
• 0
N concentration (SLN) (Coffi1or et a!., () °

1993). Cultivar Prosol 35 was grown o ~
in large pots outside in the summer in '"

o~
,I/J 3
Buenos Aires at five levels ofN fertil- <UN
ization (N ,-Ns). (a) Apparent quan- § E
:;0
tum efficiency. There is no difference .= E
a.::t
2
between the N treatments of mean rn~
value 0.05 (SE = 0.001). (b) Maximum

(I)
a:
photosynthetic rate at saturating irradi-
ance (Pmax). (c) Respiration rate at
30°C (R30). In (b) and (c), the lines are
0
fitted according to the equation Pmax or 0 2 3
R30 = b«2/ { I + exp[4(SLN .tDd)]}.tD
I». Specific leaf nitrogen (g N m- 2)
PARlm2 s (Ticha et aI., 1985). In contrast, of 0.05 mol CO 2/mol quanta (Rawson
& Constable, 1980; Connor et aI., 1993) compares well with the range 0.047 to
0.056 for C3 species (Ehleringer & Bjorkman, 1977; Evans, 1989). The compen-
sation point (at 25°C and 21 % O2) of 86 Ilmolimol air (Lloyd & Canvin, 1977) is
within the range 70 to 130 Ilmol/mol air reported for C3 species (Ticha et aI.,
1985).
The high Pmax in sunflower is of considerable interest to studies of com-
parative photosynthesis of C3 species and to crop productivity. It is achieved by
a combination of characters including, large stomatal conductance (gs), high spe-
cific rubisco activity and efficient electron transport (Delaney & Walker, 1978;
Potter & Breen, 1980; Ranty & Cavalie, 1982; Ranty et eI., 1982).
Effects of Water Deficit and Nitrogen

Net photosynthesis of plants grown in pots in glasshouses decreases with
water deficit (Boyer, 1970; Rawson, 1979; Sobrado & Turner, 1986). Rawson
(1979), for instance, showed that Pn decreased linearly with leaf water potential
('VI) at a rate of20 to 30 Ilmol CO 2/m2 s MPa in a range of 'VI from -1 to -2 MPa.
In contrast, small effects of water deficits on short-term measurements ofPn were
found in the field (English et aI., 1979; Rawson & Constable, 1980). English et
aI. (1979) suggested that osmotic adjustment may be involved in the relative
insensitivity of Pn (and gs) to water deficit in field-grown plants, a suggestion
supported by the work of Conroy et aI. (1988).
LeafN content is a major determinant of Pmax. Connor et aI. (1993) estab-
lished a common, asymptotic relationship between Pmax (range 0-40 Ilmol
CO 2/m 2 s) and specific leaf nitrogen (SLN, N content per unit leaf area) (range
0.3-2.5 g N/m2) for all leaves in plants grown at five levels of N supply (Fig.
4-5). Regardless of the cause of SLN, low supply, age, or shade-induced senes-
cence, all leaves followed the same relationship. Maximum photosynthesis was
40 Ilmol CO 2/m2 s at SLN = 2.5 g N/m2 . The same relationship explained mea-
surements made on a different cultivar by Gimenez et ai. (1994) and extended the
range of SLN up to 4 g N/m2. These analyses provide a useful integration, offer-
ing among other things the probable explanation of the previously recorded vari-
ation of Pmax with leaf position (Connor & Cawood, 1978; English et aI., 1979).
Stems and Heads

These organs make small contributions to crop photosynthesis because
their effective area and Pmax is small compared with leaves. Rawson and Con-
stable (1980) showed that Pmax of green stems was only 25% of leaves, also not-
ing that stems mostly exist in shaded environments within the crop canopy, fur-
ther restricting their contribution to crop C balance.
The C balance of heads is more complicated than that of leaves. Although
they are well-illuminated organs, heads have small photosynthetic capacity and
also large (growth) respiratory load associated with the chemical interconversions
of seed filling. There is some controversy about the importance of head photo-
synthesis to crop productivity. Rawson and Constable (1980) considered it impor-
tant, estimating a single head to be equivalent to 50 cm2 of well-illuminated leaf.
144 CONNOR & HALL
However, at that rate, even 100 000 headslha provide a negligible contribution
(0.05) to crop LAI and productivity. The measurements of Whitfield et aI. (1989)
support the conclusion that heads playa negligible role in crop photosynthesis.
They showed how the CO 2 balance of heads is negative during seed filling with
a maximum contribution of head photosynthesis equal to 30 to 45% of head res-
piration at the start of the phase.
Roots
In the field, root respiration (Rr) is measured with considerable difficulty by
difference between CO 2 efflux from soil without a root system (Rs) and soil with
roots (Rs + Rr)' Even so, Rr then includes respiration associated with microbial
activity in root exudates. Loss of C through roots by respiration, exudation, and
sloughing of root structures can comprise a significant component of a crop C
budget. Hall et aI. (1990b) have made estimates of Rr of sunflower crops in
response to temperature and water availability. Between anthesis and maturity,
daily Rr of an irrigated crop fell by 50% from an initial value of 6.6 g CO 2/m2.
Under water stress, it fell quickly to 0.37 g CO 2/m2. Accumulated Rr played a
major part in the C balance of the crop from anthesis to maturity, amounting to
167 and 110 g CO 2/m2, respectively, for irrigated and water- stressed crops, in
both cases equivalent to approximately 18% of C harvested in seed.
Canopies
The large Pmax of individual sunflower leaves ensures that crop photosyn-
thesis can increase to relatively high rates at high irradiance despite the general-
ly horizontal display of mature leaves in canopies. Measurements of crop Pn (Fig.
4-8) have confirmed this characteristic of sunflower, establishing maximum crop
rates of approximately 8 g CO 2/m2 h (ground area) at 700 to 800 W/m2 (short-
wave) leading to daily rates of assimilation of70 g CO 2/m2 (Connor et aI., 1985b;
_ 10 LAI
:.: 00
2.50
'"'e 8 0
0
a'"
u
S 6 1.40
.; 1.00
CD
.t:: 0.90
c:>-
.0
;;
4
0.43
.c:
Q.
Q.
0.27
e
u
0.21
200 400 600 800
Solar radialion (W m- 2)
Fig. 4-8. Diurnal net photosynthetic response (Pn) of well-watered sunflower crops of a range of leaf
area index (LAI) to shortwave irradiance from Connor et al. (1985b). The fitted lines are those of
the canopy photosynthesis model of Davidson and Philip (1956). In the equation: Pn = (Pmaxlk)
In[(S+K)/S exp(-kL)+K)]-LR, Pn = g CO/(m 2 ground) h, Pmax = 10.1 g CO 2/(m2Ieaf) h, k= 0.75,
S = shortwave irradiance (W/m2), K= 560 W/m2, L = LAI, and R (dark respiration) = 0.1 g CO/(m2
leaf) h.
Whitfield et aI., 1989). The exceptional photosynthetic ability of sunflower

leaves compared with other C 3 species allows it to overcome a canopy structure
that is intrinsically unsuited to high productivity. Other C3 crops with compara-
ble structure but more characteristic leaf responses, e.g., potato (Solanum tubero-
sum L.) and tobacco (Nicotiana tabacum L.), cannot match the high photosyn-
thesis rates of sunflower crops at high irradiance (Sale, 1974; Whitfield et aI.,
1980).
Water stress has counteracting effects on canopy photosynthesis. On the
one hand, smaller conductance decreases leaf photosynthetic response at all irra-
diance. On the other, wilting increases leaf angle (decreases k) and promotes a
more favorable distribution of irradiance on the canopy for photosynthesis, i.e., a
smaller proportion of leaf area is at saturating irradiance. Guiducci (1992a,b) has
documented this response, showing that the major contributor to reduced photo-
synthesis of wilting crops, the smaller interception of radiation, is substantially
offset by improved canopy light-use efficiency.
The effects of N stress on photosynthesis in young crops resemble the
effects of water stress discussed earlier, i.e., plants reduce leaf area while main-
taining photosynthetic activity per unit leaf area (Tnipani & Hall, 1996). Further,
both water and N stress affect leaf growth in similar ways, in that growth rates
are affected rather than duration of growth (Steer & Hocking, 1983; Gimenez et
aI., 1994). This supports the view that leaf expansion in sunflower is an impor-
tant adaptive mechanism to environmental stress.
Optimum Nitrogen Distribution

Leaves at the tops of canopies require large SLN to achieve large Pmax in
response to high irradiance. However, as crops grow, those leaves inevitably
become shaded as new leaves expand above them. Then, under low irradiance,
they benefit less from large SLN because their photosynthetic performance is
more related to £ (see Eq. [1]) which is constant over a wide range of SLN
(Connor et aI., 1993). The question then arises if the vertical distributions ofSLN
established by mobilization of N from mature leaves to expanding leaves and
from senescing leaves to seed are consistent with maximum canopy photosyn-
thesis. Stated another way, the question asks, would different mobilization
responses establish optimum profiles that would maximize photosynthetic pro-
ductivity with limited supply ofN?
There is no single optimal profile because performance of each LAI and
SLN combination depends upon irradiance. The largest gains, compared with say
an even vertical profile, occur in dense rather than open canopies, under high
rather than low irradiance, and under small rather than large SLN (Hirose &
Werger, 1987). Analyses with sunflower suggest that neither the vertical profiles
of N established during the vegetative (Gimenez et aI., 1994) nor during the
reproductive phase (Sadras et aI., 1993a) are optimal (Fig. 4-9). In the case of
vegetative canopies, the productive loss was significant, being of the order of
14% in the extreme cases established by Gimenez et ai. (1994). That study
showed that the effects of N supply on productivity extend well beyond their
effect on photosynthesis so that resolution of this issue of optimal crop behavior
146 CONNOR & HALL
30
(8)
25
20
iii
.0
E
:::J 15
c:
1a
~ 10
DAS - 42
LAI - 1.30
5 TNC _ 2.67 g m- 2
0
1 2 4 5 6
Specific leaf nitrogen (g N m-2)
30
(b)
25
20
.0
iii
E
:::J
c: 15
1aCD
-l
10
DAS _ 56
LAI _ 3.32
5 TNC _ 6.00 g m- 2
0
0 3 4 5 6
Specific leaf nitrogen (g N m- 2)
30
Fig. 4-9. Profiles of observed (open sym- 25

bols) and optimal (closed symbols)
specific leaf N concentration (g N/m2)
20
of sunflower crops (Sungro 385) of
defined leaf area index (LAI) on three iii
.0
occasions, at bud visible (42 dafter E
:::J 15
c:
sowing, DAS), at midstem extension
1a
(DAS 56), and at first anthesis (DAS CD
-l 10
71) at Cordoba, Spain. Crops were
sown at 5.7 m-2 and received 25 g DAS -71
N/m 2 at sowing. The optimal distribu- 5 LAI - 3.21
tions are those of the same total N con- TNC _ 7.83 g m- 2
tent (TNC) that would maximize crop 0

daytime photosynthesis at daily short- 0 2 3 4 5 6
wave radiation of 25 MJ/m2 (after
Gimenez et ai., 1994). Specific leaf nitrogen (g N m- 2)
for maximum productivity will increasingly depend upon the development of

techniques to study the integration of physiological responses that is presented in
a subsequent section.
BIOMASS PARTITIONING
Assimilate formed in photosynthesis is the major component of plant bio-

mass. It is exported, as sucrose from leaves, or intermediate storage organs such
as stems, to other organs where it is built into new biomass or respired to support
their metabolism. The amount of assimilate required per unit growth of biomass
varies between organs, with ontogeny, and with environmental conditions. These
differences are largely a result of the growth respiration cost of forming biomass
of differing chemical composition, but the impact of the potentially important
maintenance component has yet to be satisfactorily resolved and has only rarely
been estimated (e.g., Hall et aI., 1990b). The proportion of C per unit biomass of
each organ (Fig. 4-10) reflects the outcome of these interactions and the mineral
content. Progress has been reported in understanding the processes, pathways,
and control mechanisms involved in the partitioning of newly formed assimilate
and its transport from the leaves to intermediate storage sites and final destina-
tions (Gifford et aI., 1984; Wardlaw, 1990), as well as in the efficiency with which
it is transformed into new biomass (Penning de Vries et aI., 1983; Loomis &
Connor, 1992).
The result of assimilate partitioning can be assessed with partitioning coef-
ficients (PC), which describe the proportion of the total biomass increment parti-
tioned to specified organs. However, the nature of biomass and the processes
involved are better assessed by correcting for its production value (PV cb), rather
Ic
a
t
0
0.7
O.S
---
-0--
--0---
---.--
fine roots
stem plus taproot
leaf
receptacle
chaff
seed·W
t(::
.'.--.
~:::_<>-.-o
c. ---<)0-- seed·D
a.
0
0.5
~
c
0
.0
iii
(,)
0.4
c
'"
0>
0 fN FA PM
0.3
40
j
SO
'" 80 100
.'M
120
Days after sowing
Fig. 4-10. Changes with time and phenological development (BV = bud visible. FA = first anthesis,
and PM = physiological maturity) in the organ C proportion of sunflower. Values for seed refer to
crops irrigated throughout (seed-W) or droughted 80 d after sowing (seed-D), values for remaining
organs did not differ between treatments. Mineral [i.e., (ash X 0 6. ), cf., Penning de Vries et aI.,
1983) content differed between organs, being approximately 110 g/kg in leaves, approximately 60
g/kg in receptacle and approximately 15 g/kg in seed, and increased during grain filling in nonseed
organs. Measurement techniques and crop management described in Hall et al. (1990b).
148 CONNOR & HALL
than its mass, to account for differences in growth respiration of organs of differ-
ent elemental composition (e.g., Hall et aI., 1989; Sadras & Connor, 1991). The
most complete description would include the effects of maintenance respiration,
but this is technically difficult and is rarely attempted. The distinction between
partitioning based on biomass or PV cb is important in the following discussion,
which reviews the present status of attempts to describe assimilate partitioning
between organs of sunflower crops in response to ontogeny, environment, and
management.
Pre-Anthesis
Horie (1977) pioneered attempts to describe assimilate partitioning, in

terms of PC, in sunflower crops, using growth analysis, estimates of organ
growth, maintenance respiration, and responses to temperature. The pattern for
the preanthesis period was characterized by stable, small (approximately 12%)
PC for roots, early dominance of partitioning to leaves that was later transferred
to stems, and increasing importance of the head in the 10-d period preceding
anthesis. A feature of his analysis is the continuous change in PC with ontogeny.
Trapani et al. (1994) showed, using a slightly different approach, that constant
values for organ PC for each of three defined phenophases (Table 4-1) provided
an effective description of biomass partitioning between organs over a range of
crops and cultivars grown without water and N stress. A notable feature of both
analyses is the dominance of the stem for much of the preanthesis period, with
peak PC of approximately 0.60 for periods of close to 30 d. The partitioning pat-
terns based on PVcb were consistent with those based on biomass, the main dif-
ference being the greater values for lamina (in consequence, lower values for the
other organs), reflecting the impact of lamina respiration (Horie, 1977).
The estimates of pre anthes is biomass partitioning reported by Horie (1977)
and Trapani et al. (1994) were derived from measurements made on crops well
provided with water and nutrients. It is important for practical and theoretical rea-
sons to establish what effects environment, cultivar, and management may have
on partitioning. Villalobos et al. (1996) have shown that increasing density from
0.5 to 10 plants/m2 decreased partitioning to leaves and increased that to stems
and head. Interestingly, changes in partitioning over the range 5 to 10 plants/m2
Table 4-1. Values of partitioning coefficients (PC) (means ± SE) calculated for the phases emergence
to bud visible (E-BV), bud visible to start of rapid head growth (BV -SHRG), and start of rapid head
growth to eight-row anthesis (SHRG-A). Values are means of three experiments involving five
genotypes (lines and hybrids). Start of rapid head growth occurred 14 d before anthesis. Sum of
organ PC for each period does not necessarily equal 1.00 due to the technique used for estimation
(data from Trapani et aI. , 1994).
Period
Organ E-BV BV-SRHG SRHG-A
Stem 0.36 ± 0.02 0.60 ± 0.03 0.60 ± 0.03

Laminae 0.42 ± 0 0. 3 0.29 ± 0 .03 0.15 ± 0 0. 2
Fine roots 0.19 ± 0 0. 3 0.08 ± 0.01 0.05 ± 0 0. 2
Head 0.03 ± 0 0. 1 0.16 ± 0.03
(usual commercial densities) were not significant, but greater partitioning to stem
has been recorded at 15 compared with 5 plants/m2 (Steer et aI., 1986; Merrien et
aI., 1983). These responses to density contrast with those induced by shading,
which decreased partitioning to the capitulum (Villalobos et aI., 1992; Sinsawat
& Steer, 1993). The contrast between the responses to these two factors suggests
that density effects are not simply mediated by assimilate availability, and that
other controls such as light quality (Sanchez et aI., 1994) may be involved.
Sinsawat and Steer (1993) also found that although shading substantially reduced
the fraction of total biomass in the capitulum, it had no effect on the distribution
of biomass between receptacle, involucral bracts, and florets. Evidently, the con-
trols over interorgan partitioning differ from those over intraorgan partitioning.
Elucidation of the mechanisms controlling partitioning to the capitulum is partic-
ularly important given its relation to seed number and yield.
Information on the effects of other stresses on partitioning of biomass is
sparse. Exposure of plants to N levels ranging from severely inadequate to ade-
quate between FI to FA had no effect on the partitioning of biomass between
aboveground organs (Sinsawat & Steer, 1993) while exposure to preanthesis
water stress reduced the assimilate flux to the head much less than to the stem
(Sadras et aI., 1992). Sobrado and Turner (1986) reported similar, although less
marked, responses. Interpretation of these patterns of partitioning between shoot
organs becomes complex because increases in root/shoot ratio (RlS) are an addi-
tional expected outcome of exposure to N and water stresses (Loomis & Connor,
1992). Sadras (1990) demonstrated consistent increases in RlS of three cultivars
to preanthesis water stress, but other workers have reported no change or the
opposite result (Sobrado & Turner, 1986; Lovett & Campbell, 1973). Connor et
aI. (1985a) reported increases in the root length! leaf area ratio (RLILA) in
response to water stress. Variation in RL/LA does not necessarily reflect changes
in biomass partitioning, but these changes are consistent with the expected
improvement in root functioning relative to shoot.
Further research on these issues is clearly needed, but it seems likely in
sunflower, as in other species, that exposure to water and N stresses before anthe-
sis (i.e., before the seed becomes the dominant sink) will produce changes that
favor partitioning to the root, consistent with Brouwer's (1962) concept of func-
tional equilibrium between root and shoot. Responses of sunflower RlS to root
anoxia (Kriedemann et aI., 1983) and root temperature (Szaniawski, 1983) sup-
port this argument.
There is little information on cultivar effects on biomass partitioning.
Sadras (1990) showed that RlS ratios of standard height, semidwarf, and dwarf
cultivars increased with degree of dwarfism and that cultivar ranking was main-
tained between BV to FA despite a continuing fall in RlS.
Post-Anthesis
After anthesis, first heads, and then seed, become the dominant sink for
assimilate. Seed growth is sustained by both current photosynthesis and mobi-
lization of stored assimilate which results in substantial net loss of biomass from
150 CONNOR & HALL
nonseed organs between LA and PM (Hall et ai., 1985; Connor et ai., 1985a; Hall
et ai., 1989; Iaafar et ai., 1993). The use of information on organ dry weight
dynamics and respiration alone to estimate partitioning of gross assimilate
(Horie, 1977) is not applicable to circumstances in which an important part of the
dry weight increase of an organ does not derive from current photosynthesis, and
other techniques such as crop and organ C balances, or labeled C dynamics, must
be used. A C-balance analysis showed that assimilate fixed before anthesis and
stored mostly in the stem and receptacle contributed an important proportion (15
and 27% in irrigated and stressed crops, respectively) of the total assimilate (i.e.,
fixed plus respired) flux to the seed (Hall et ai., 1990b). Somewhat greater esti-
mates were obtained using the less-accurate, labeled-C technique (Hall et ai.,
1989). Absolute and relative contributions of stored preanthesis assimilate to seed
filling in nonstressed sunflower were roughly double those measured by similar
techniques in cereals of similar aboveground biomass at anthesis.
The biomass formed during preanthesis that is mobilized to support seed
growth includes both carbohydrates and nitrogenous compounds (Steer et ai.,
1985b; Hall et ai., 1989), but the former are dominant. Levels of water soluble
carbohydrates (WSC, largely fructose and fructans) can reach high values (43%
in stem, 67% in receptacle), and the dynamics of WSC suggest that storage of
these substances in the receptacle increases for some time after anthesis.
However, about 30% of WSC present in the stems and receptacles at anthesis
remained at PM (Hall et ai., 1989). The importance of the contribution of stored
assimilate to seed filling in sunflower emphasizes the need for further research.
It is important to define the dynamics of assimilate partitioning between struc-
tural and nonstructural components of the stem and receptacle in order to define
their maximum storage capacity, and to examine the possible role of the recepta-
cle as an intermediate storage organ.
Source-sink relationships during seed filling may affect the relative contri-
bution of preanthesis assimilate to seed filling. Sadras et ai. (1992) have estimat-
ed proportional contributions to seed filling between 0.1 and 1.0 depending on the
ratio between seed number and assimilate produced after anthesis in standard
height (but not dwarf) cultivars. Schneiter et ai. (1987) reported greater stem
breakage in sunflower crops subjected to heavy defoliation, suggesting there may
be some linkage between the levels of assimilate in the stem and susceptibility to
breakage. In maize, low concentration of stem solutes has been linked to suscep-
tibility to stalk rots (e.g., Dodd, 1980). If this linkage is operative in sunflower, it
may impose a ceiling to the selection of cultivars with greater mobilization of
stem-stored assimilate.
Together with crop biomass at anthesis and postanthesis crop photosynthe-
sis, partitioning of assimilate to seed is one of the important determinants of har-
vest index (HI). Recent experiments and analyses have generated useful insights
into how cultivar attributes (phenology and dwarfism), management (density),
and environment-management interactions may affect this important ratio. Sadras
and Connor (1991) demonstrated that HIpv (i.e., corrected for synthesis costs)
increased curvilinearily from 0.28 to 0.62 (0.13-0.46 for HI) with changes in the
fraction of water transpired after anthesis by the crop, normalized for changes in
evaporative demand between pre- and postanthesis, from 0.04 to 0.41. They also
showed how this response may be related to the potential HI and the potential
contribution of preanthesis assimilate to seed. The latter attributes are genotype
dependent and affected by phenology and plant stature.
Analyses using a simulation model by Sadras and Villalobos (1994) under-
line the importance of the interactions between phenology and water availability
operating on HI and crop biomass. Lengthening E to FA decreases HI and yield
under water-limited conditions, and increasing the duration of seed filling with
greater water availability increased both HI and yield. Genotype and density
effects on HI are illustrated by the results of Villalobos et al. (1996). In crops
grown over a range of densities from 0.5 to 10 plants/m2, HI of the shortest-cycle
hybrid increased over the whole range, while that of the longer-cycle hybrids
reached a plateau between 5 and 10 plants/m2 • This difference was attributable to
the limited plasticity in seed number and weight of the shortest-cycle hybrid. This
raises the important point that HI of sunflower plants at low densities may be dif-
ferent to those at commercial planting rates. This contrasts with tillering in cere-
als in which equilibrium between nonseed and seed biomass may be maintained
over a wide range of densities.
These observations lead to the proposition that even under conditions of
unlimited resources there is an optimum combination of pre- and postanthesis
duration and density that maximizes HI and/or yield. Simulation analyses of the
effects of extending the preanthesis phase from 82 to 104 d suggested yield would
increase from 1.3 to 5.5 tiha, while peak HI was achieved with a preanthesis dura-
tion of94 d (Sadras & Villalobos, 1994). The increase in yield included compo-
nents attributable to greater radiation as the season progressed and a longer peri-
od of full interception of radiation. Clearly, genotype X environment interactions
upon partitioning, HI, and yield can be complex. Experimentation and simulation
analyses can both contribute useful insights into these interactions.
YIELD FORMATION AND REALIZATION
Components of Yield
Seed number and size together with oil content and type are the principal
components of yield in sunflower. There also is some interest in protein content
(e.g., Stanojevic et aI., 1992). Seed number, size, and oil content are determined
sequentially during the life cycle of the crop, with some overlapping between
contiguous phases. Some progress has been made in determining the phenophas-
es when each component is affected by environmental factors. Two experimental
approaches have been especially useful. One applies a series of brief stresses, the
other employs manipulations (e.g., density, fertilization, and early defoliation)
that affect crop performance over longer periods. The approaches are comple-
mentary and have revealed important interactions between the crop and environ-
ment. In the following, most emphasis is given to data obtained in experiments
conducted within a clearly established phenological framework. This applies
especially before the BV stage and during seed filling.
152 CONNOR & HALL
Seed Number
Seed number is the yield component with the largest phenotypic and geno-
typic variation (e.g., Steer et aI., 1985a; Steer & Hocking, 1987) and is the one
most affected by vegetative manipulations such as defoliation (Schneiter et aI.,
1987). In sunflower, seed number is determined between FI and the start of rapid
kernel growth, a long period compared with other crop species such as wheat and
maize. This has been shown in experiments involving brief exposures to shading
(Chimenti & Hall, 1992) or water stress (Talha & Osman, 1975; Yegappan et aI.,
1982; Hall et aI., 1985) and also from defoliation which continues to exert effects
after FA (e.g., Cardinali et aI., 1982; Schneiter et aI., 1987). The processes
involved in determining final seed number include floret differentiation, floret
growth up to anthesis, fertilization, and embryo abortion.
The upper limit to seed number per plant is set by the number of florets per
head that are differentiated between FS 5 (Marc & Palmer, 1981) when the pri-
mordia of sterile ray florets appear at the periphery of the floral disc, and FS 8,
when floret primordia cover the entire disc. This may last 7 to 12 d under field
conditions, but longer durations have been observed in controlled environments
(Palmer & Steer, 1985). The number of primordia on the floral disc depends upon
the number of floret rows (parastichies) initiated, their length, and the degree of
amalgamation of rows that occurs towards the center of the disc. A wide range
(700-1900 florets/head) of phenotypic and genotypic variation was shown to be
associated with the duration (the integral across time) of the generative area, an
inner disc on the receptacle limited by the front of floret initiation (Palmer &
Steer, 1985). Both genotype and N availability prior to floret differentiation may
affect the size of the generative area, and hence the number of florets established
per head (Steer et aI., 1985a; Palmer & Steer, 1985). Low N availability reduced
receptacle area, generative area, and floret number to about one-third of normal
values. Much of these effects appear to derive from a reduction in the duration of
FS 1.3 to FS 8, largely through delay in the achievement ofFS 1.3, although the
rate of floret differentiation also may be reduced (Hocking & Steer, 1989).
Effects of water stress on floret initiation also have been found in pot exper-
iments (Marc & Palmer, 1976), but the relevance of these results for field condi-
tions has yet to be established. Clearly, receptacle size at FS 5 and its expansion
up to FS 8 set an upper limit to the maximum number of florets. Mean area per
floret (range 0.045-0.053 mm2) at FS 8 varied little between genotypes over a
twofold range in floret number, suggesting that average floret size at FS 8 may be
fairly conservative (Palmer & Steer, 1985). It should be remembered, however,
that at any stage between FS 5 and FS 8, florets initiated at the periphery are the
oldest and largest on the disc and develop first throughout the remainder of the
crop cycle. Labeled metabolites are preferentially partitioned to peripheral
regions of the developing capitulum (Hernandez & Palmer, 1992). During seed
filling, dry weight and lipid accumulation gradually transfer from outer to inner
seed (Goffner et aI., 1988).
There has been considerable interest in the causes and control of floret dif-
ferentiation (Hernandez & Palmer, 1988; Hernandez & Green, 1993) as well as
some preliminary investigation into the role of plant growth regulators in this
process and their effects on yield (Hernandez, 1992). Some genotypic variation
in florets per row has been found (Palmer & Steer, 1985) but was too small to
compensate for differences in number of rows. Excision of upper leaves and
involucral bract primordia at FS3 increased the number of floret primordia pre-
sent at FS8 and seed yield per plant, as well as increasing the rate of progress of
floral differentiation (Hernandez, 1993). This suggests that competition for
assimilates between the growing capitulum and the upper leaves and bracts (or
some related hormonal signal) may affect potential floret number.
Once floret primordia have differentiated, they continue through an order-
ly process of development and growth that culminates in anthesis. Exposure to N,
shade, or water stress during this period reduces final seed number per head
(Yegappan et al., 1982; Steer et aI., 1984; Hall et al., 1985; Chimenti & Hall,
1992). There also is evidence that seed number is positively correlated with radi-
ation and negatively with temperature during the FI to FA phase (Rawson et al.,
1984), a response similar to that found in wheat (Fischer, 1985). The effects of
stresses are most clearly seen in smaller fertility ratios (seed produced/florets
available) in the central portion of the capitulum (the "empty center" syndrome).
Empty centers, that can be a problem even in commercial crops well provided
with water and nutrients, may have a genotypic component that is often seen in
dwarf cultivars. The physiological basis of this effect remains uncertain.
Mc William and English (1978) found a general relationship between inflores-
cence diameter and seed number per inflorescence, but it is not known how much
variation in the number of florets initiated and the fertility ratio contributed to the
final result.
Sinsawat and Steer (1993) have shown that shading and restricted N supply
between FS 8 and anthesis reduces the growth of the capitulum, leading to small-
er individual floret cross-sectional area at anthesis at all positions on the recepta-
cle. Variations in floret height were relatively small. The data of Villalobos et al.
(1996) show a generalized, if not always consistent, negative relationship
between the apparent density of reproductive structures and fertility ratio across
the outer, middle, and central portions of the capitulum in crops grown over a
range of densities. These observations are consistent with the hypothesis that lack
of seed from the central florets is due to increased competition for space on the
capitulum. Poor or delayed vascularization of the central portion of the capitulum
also may be involved (Durrieu et al., 1985). However, these are not the only alter-
natives.
In wheat, part of the floret popUlation lose competence before anthesis, a
loss that can be exacerbated by shading (e.g., Stockman et aI., 1983) and expo-
sure to water that stress affects pollen formation (Morgan, 1980). Whether simi-
lar processes occur in sunflower has not been evaluated. Restriction of pollina-
tion to the central florets on the capitulum has shown they are capable of pro-
ducing seed of normal size, suggesting their competence and capitulum vascular-
ization is not affected by position (Steer et aI., 1988). However, information on
the response of the fertility ratio of the central florets to the relief from competi-
tion is needed before the loss-of-competence hypothesis can be discarded with
confidence. Dominance effects, presumably mediated by hormonal signals rather
than availability of space, of early established seed or fruits over later-established
154 CONNOR & HALL
ones, are known to occur in other species (Freier et aI., 1984; Bangerth, 1989) and
are increased by exposure to stress. These processes also may operate in sun-
flower.
Separate, albeit potentially important causes of small seed number per
head, are self incompatibility (Luciano et aI., 1965), poor pollination (Roath &
Miller, 1982), and B deficiency (B1amey et aI., 1997). In those cases, florets that
fail to set seed tend to be distributed randomly across the head rather than con-
centrated at the center of the capitulum as "empty centers." Considerable
progress has been made in reducing self-incompatibility and improving self-pol-
lination in modem oil seed hybrids (e.g., Vranceanu et aI., 1988), but advantages
still accrue from using bees (Apis melli/era L.) as pollinators (Virupakshappa et
aI., 1992), suggesting room for further improvement. Advances in understanding
the molecular biology of self-incompatibility may contribute to this improve-
ment.
Brief exposures to drought or shade after anthesis increase the proportion
of empty hulls ("flats") (Yegappan et aI., 1982; Hall et aI., 1985; Chimenti &
Hall, 1992) showing that abortion may reduce seed number during this
phenophase. The detailed physiology of these effects has not been studied and
would benefit from the kind of attention paid to seed loss in maize in response to
water stress (e.g., Schussler & Westgate, 1991; Bassetti & Westgate, 1993). The
final period for seed number adjustment in sunflower appears to include the early
stages of embryo growth (Chimenti & Hall, 1992).
Focused exposure to brief stresses is the effective technique for identifying
the periods of sensitivity and the processes involved. The impact of density is dif-
ferent, however, because density stress operates throughout most of the crop
cycle. There are few experiments in which the effect of density on the actual
number of florets (as distinct from the sum of filled seed plus flats) has been fol-
lowed. The results of Steer et aI. (1986) suggest that an increase in density from
5 to 15 plants/m 2 led to a 12% reduction in floret numberlhead and a decrease in
fertility ratio from 0.68 to 0.38. The long-season hybrids used by Villalobos et aI.
(1994), which reached yield plateau at fairly low populations, showed a drop in
apparent fertility ratio (estimated from seed plus flats) from approximately 0.9 at
2 plants/m2 to between 0.58 and 0.78 at 10 plants/m 2.
It has been argued, largely on the basis of plant responses to severe N defi-
ciency, that seed number in sunflower is determined during floral initiation (e.g.,
Steer et aI., 1984, 1986). The previous discussion shows, rather, that seed num-
ber is the result of a ceiling value fixed during floret initiation, then subjected to
a series of adjustments up to the start of embryo growth. Fertility ratios seldom
exceed 0.7 to 0.8 in crops growing at commercial densities that are well provid-
ed with water and nutrients (e.g., Steer et aI., 1986; Hall et aI., 1985; for an excep-
tion see Palmer & Steer, 1985), and even in plants growing at very low density
(Villalobos et aI., 1994). This suggests that sunflower, as other crop species, pro-
duces more florets per capitulum during initiation than are likely to set seed, and
that loss of florets and fertilized ovules is part of a normal adjustment to envi-
ronmental conditions during the crop cycle. In this way, the ceiling number estab-
lished at floret differentiation is unlikely to present a barrier to seed-number
determination in commercial production, although it may limit seed number of
crops at extremely low densities in very short-season cultivars (Villalobos et ai.,

1994). Future work should focus on the causes and physiological mechanisms of
reduction in seed number immediately prior to, during, and after anthesis.
Seed Weight
Sunflower seed comprise pericarp, derived from the ovary wall, the true
seedcoat, and the kernel which is mostly embryo (Knowles, 1978). During
growth, sucrose and nitrogenous compounds pass from the apoplast of the seed
coat into the growing embryo (Wolswinkel, 1987). Abscisic acid (C2oH2004) con-
tent of the kernel increases during its growth (Karyagina et ai., 1988) indicating
that it may playa role in the control of assimilate uptake (Thome, 1985) although
other mechanisms have been proposed (Wolswinkel, 1987). The pericarp (hull)
weighs about 2 mg at anthesis, grows for around 8 d before the embryo starts to
grow rapidly. Hull growth ceases when the embryo is about one third of its final
weight (Fig. 4-11) (Villalobos et ai., 1996). One report (Karyagina et ai., 1988)
suggests a temporary reduction in growth rate of the whole seed associated with
the switch in growth from peri carp to kernei. A decrease in indole acetic acid
(C IOH 9N0 2, IAA) content of seed is associated with this switch in growth pattern.
Interestingly, the pericarp appears to be the last part of the seed to lose water dur-
ing maturation (Karyagina et aI., 1988).
The pericarp comprises between about 20 (modem oilseed hybrids) and
40% (confectionery types) of final seed weight, a greater proportion than for
equivalent structures in other seed crops. The differences in pattern and timing of
growth of pericarp and kernel have important implications for crop response to
stress and for seed quality. Although growth of seed in the inner part of the capit-
ulum is initiated later than at the periphery (Unger & Thompson, 1982; Goffner
et aI., 1988), seed in all positions appear to reach PM close to the same time from
120
...
100
Ci 80
E. o
E Kerne!
Ol 60
·m
il:
~ 40
0
20 Hull
0
5 10 15 20 25 30 35 40 45
Days after first anthesis
Fig. 4-11. Patterns of growth of the whole seed, hull, and kernel of Arbung E353 growing in the three
outer rows of the capitulum. Sloping straight lines show regressions fitted to the linear-growth
phases, horizontal lines to the plateau of maximum weight. Full symbols are observed values,
empty symbols are estimates obtained assuming constant hull weight at 16 d after first anthesis (F.
Orgaz, unpublished data).
156 CONNOR & HALL
FA (Unger & Thompson, 1982; Hall et aI., 1989). The rate as well as the duration
of seed filling also may be less for the inner seed (Villalobos et ai., 1994). The
consequence is smaller seed at the center of the capitulum. Newcomb (1973) has
studied the structural and ultrastructural aspects of the early stages of embryo
development in sunflower, but there is little information of the processes involved
and their implications for seed size and quality.
Large genotypic and phenotypic variations in seed size have been reported
(e.g., Palmer & Steer, 1985; Steer & Hocking, 1987; Hall et aI., 1985; Villalobos
et aI., 1994) and have been related to rate of head expansion close to flowering
(Steer & Hocking, 1987). These differences appear to relate more to variations in
seed or embryo growth rate from LA to PM than to changes in duration of seed
growth (Hall et aI., 1985; Villalobos et ai., 1994). Thus, the difference in seed size
(approximately 27%) between 'Arbung E353' (large seed) and Sungro 385 (small
seed) that was maintained over the range of crop densities from 0.5 to 10
plants/m2 was associated with a slightly larger difference in embryo growth rate.
Duration of seed filling between extreme genotypes varied between 39 and 46 d
while mean duration for Sungro 385 was 5% greater than Arbung E353
(Villalobos et ai., 1994). There is little information on genotypic variation in seed
growth in sunflower and none on the effects of temperature, which is known to
affect both rate and duration of seed filling in other crops (e.g., Chowdhury &
Wardlaw, 1978).
Nitrogen, water, and density stresses affect seed size. Variations in supply
imposed at different stages of plant growth have shown that N supply effects seed
size particularly during FI to FA, with minor effects during later stages (Steer et
aI., 1984). In contrast, brief exposures to water stress reduced seed size at all
stages between FS 6 and mid-seed filling (Talha & Osman, 1975; Yegappan et ai.,
1982; Hall et aI., 1985). Intense defoliation had substantial effects on seed size
immediately prior to and after anthesis (Schneiter et ai., 1987). Reduction of
interseed competition by restriction of pollination allows normally small seed
close to the center of the capitulum to grow larger than on fully populated heads
(Steer et aI., 1988). It would be useful to establish whether the effects of pre an-
thesis stress reflect limitations of space on the receptacle or within small peri-
carps. Steer et aI. (1988) found that partial defoliation at anthesis reduced hull
more than kernel weight. Taken together, these results suggest that although seed
retain an important capacity to respond to improved conditions after anthesis,
restriction of seed size imposed by stresses before anthesis are likely to be trans-
lated into smaller seed at harvest.
Stress during seed filling is likely to reduce individual seed weight in most
circumstances. Nitrogen stress after anthesis has small impact, a not unexpected
result, given the over-riding importance of stored N to seed protein in this crop.
The effects of long-term stresses, such as defoliation after anthesis (Steer et aI.,
1988; Schneiter et ai., 1987; Cardinali et ai., 1982), or density (Villalobos et ai.,
1994), on seed size are consistent with the picture emerging from brief exposures
to stress. Stress imposed by density is active throughout most of the crop cycle,
presumably impacting on both the preanthesis growth of florets and on seed fill-
ing. Defoliation, when intense and applied close to anthesis, usually restricts seed
filling. In both cases, poorly understood mechanisms of yield compensation
between seed size and number come into play. There is some evidence that plant
growth regulators playa role in the control of seed size (Karyagina et ai., 1988).
Seed Number/Size Interactions
The overlap between the periods when seed number and seed size are influ-
enced by environment complicates analysis of the interactions between the two
components. This contrasts with other crop species, such as wheat (Fischer, 1985)
and, to a lesser extent, maize (Fischer & Palmer, 1983), in which the responses
are more easily separated. Density effects on both seed size and seed number took
place over the range 0.5 to 10 plants/m2 explored by Villalobos et ai. (1994). This
also was true for the range 5 to 10 plants/m2 in which the long- cycle hybrids
reached their yield plateau, with increments in seed number/square meter, be-
tween 32 and 49% according to genotype, being balanced by proportional reduc-
tions in seed size. This capacity for yield adjustment through both components
over a wide range of conditions underlines the need for better understanding of
the physiological mechanisms underlying these responses.
Seed Oil Content
Small quantities of lipids (10-30 glkg) are normally found in all tissues of
sunflower, much of it associated with cellular and subcellular membranes. Seed
is no exception during its early stages of growth (Harris et ai., 1978). Rapid depo-
sition of reserve triacylglycerols (TAG) which form the greater part ofthe oil con-
tent of seed, only begins some days after the start of rapid embryo growth
(Villalobos et aI., 1996), and little oil is deposited during the first third of the
seed-filling period (Harris et aI., 1978). This delay between anthesis and the start
of oil deposition has been found in other species also (Murphy, 1990; Tzen et aI.,
1993). The sharp change in partitioning oflabeled oleate between polar lipids and
TAG at approximately 10 dafter anthesis found in sunflower by Garces and
Mancha (1989) is another manifestation of these ontogenetic changes. At matu-
rity, almost all the oil present in seed is located in the kernel (McWilliam &
English, 1978), largely within oil bodies. The hull contains about 20 to 30 glkg,
explaining the level of wastage if seed is dehulled before crushing. There has
been considerable progress in recent years in understanding glycerolipid synthe-
sis (Browse & Somerville, 1991) and the nature and formation of oil bodies
(Huang, 1992). Much of the new information relates to species other than sun-
flower and to organs other than seed.
Synthesis of storage lipids is complex involving metabolic transformations
in the cytosol, proplastids, and endoplasmic reticulum of the embryo cells
(Browse & Somerville, 1991). Dihydroxyacetone phosphate derived from gly-
colysis in the cytosol is converted to glycerol-phosphate, the source of the glyc-
erol skeleton of TAG, in the cytosol, or it may move across the double mem-
branes of the proplastid envelope. There, after further transformations, acetyl
CoA and malonyl CoA are formed. These are the primer and building blocks,
respectively, for a stepwise elongation of the fatty acid chain, a process involving
a multi enzyme fatty acid synthase (FAS) complex and a low molecular weight
158 CONNOR & HALL
acyl carrier protein (ACP). At each turn of the FAS cycle, a two-C moiety is
attached to the growing chain until a saturated 16-C fatty acid (CI6:0, palmitate)-
ACP complex is produced. Further elongation (not mediated by FAS) to 18:0
(stearate)-ACP and an initial de saturation to 18:1 (oleate)-ACP may occur with-
in the proplastid. Separation of ACP from the fatty acid may take place at any
stage after the formation of the 16:0-ACP complex, but in sunflower most fatty
acid molecules presumably progress to the 18:l-ACP stage before the complex is
cleaved.
Free fatty acids move through the double membrane of the proplastid and
are converted to acyl-CoA at the outer membrane, before entering the cytosol.
The putative pathway for synthesis of sunflower TAG continues on the rough
endoplasmic reticulum, where it is assembled in a series of steps from cytosolic
pools of glycerol phosphate and acyl-CoA. This process is not fully understood,
but it seems clear, by analogy with other species, that the acyl moieties are added
to the glycerol skeleton one at a time, and that further de saturation to 18:2 (linole-
ic) fatty acid may occur during TAG assembly. The relative importance of phos-
phatidyl choline and phosphatidic acid as steps in the pathway of diacylglycerol
synthesis is an important issue that awaits resolution (Browse & Somerville,
1991). Successful elucidation of the subcellular localization, the details of the
biosynthetic pathways, and of the regulation of the activity of the key enzymes in
the TAG pathway probably hold the key to an effective and systematic applica-
tion of biotechnology to the manipulation of sunflower oil quality (Knauf, 1987;
Sommerville & Browse, 1991; Ohlrogge et aI., 1991).
Huang (1992) and Tzen et al. (1993) propose that oil bodies are globes of
TAG covered with small amounts of phospholipid «1 % in maize) and a specific
oil-body protein (oleosin) «1.5% in maize) that form a half-unit membrane. The
oleosin has a long hydrophobic stalk, attached to an outward-facing hydrophylic
cap, that penetrates into the TAG region, an arrangement that contributes to oil
body stability. Oil bodies are formed by deposition of TAG between leaflets of
the endoplasmic reticulum, and this structure buds off to become an oil body.
Oleosins are synthesized on mitochochonrial ribonucleic acid (mRNA) associat-
ed with the endoplasmic reticulum and are apparently inserted into the half mem-
brane at about the time of budding, although there is controversy about this point
(Murphy, 1990). Confirmation of this description and the improved understand-
ing of the control of oil-body formation (size, number, and timing) has impor-
tance for sunflower which is presumably comparable to that of starch bodies for
cereal seed physiology (e.g., Martinez Carrasco et aI., 1988). The combination of
oleos ins and phospholipids on the surface of the oil bodies ensures that they do
not coalesce under extreme treatments or in air-dried seed.
The pattern of protein deposition in seed contrasts that of oil, proceeding in
concert with seed growth so that the concentration of protein in seed dry matter
remains fairly constant over time (Goffner et aI., 1988.). This et aI. (1988) have
characterized the dynamics of accumulation of the storage proteins helianthin and
albumin. Because oleosins are obligate components of oil bodies, a part of the
total protein content is associated with oil deposition. Studies have shown little
variation in seed oil concentration at maturity across positions in the head of non-
stressed crops (McWilliam & English, 1978; Goffner et aI., 1988), even though
the inner seed commence growth and oil deposition later than those towards the
periphery. The differences in temporal patterns of deposition of oil, protein, and
other components of seed may have profound effects on oil content and concen-
tration of bulk seed, particularly from crops subject to stress during seed filling.
Genotype and environmental factors, including temperature, water stress,
and N have been shown to affect the amount and concentration of oil in seed.
Genotype effects are largely attributable to the final proportion of peri carp to ker-
nel (McWilliam & English, 1978) but deviations from that general relationship
have now been identified (Mantese & Medan, personal communication), indicat-
ing the existence of further control over kernel oil content. Seed oil content was
negatively related with temperature during seed filling in controlled-environment
studies (Harris et ai., 1978; but cf. Canvin, 1965; Rawson et ai., 1984) and in
time-of-sowing experiments (Seiler, 1986b). The cause of these reductions is
uncertain. A greater proportion of pericarp, due to shortening of the seed-filling
period at high temperature, can only explain part of the effects found by Harris et
ai. (1978). Re-evaluation of earlier reports (e.g., Johnson & Jellum, 1972; Unger,
1980; Unger & Thompson, 1982) of seed maturing late in the season at low tem-
perature shows that the low oil content ofthose seed (i.e., reflecting a supposed-
ly positive effect of temperature on oil content) was usually associated with
small or light seed, possibly due to problems with pollination or subsequent seed
growth. The issue deserves further study.
Exposure to brief periods of water stress during seed filling reduces oil con-
tent in association with increases in the proportion of hull (Hall et ai., 1985,
1989). Talha and Osman (1975) reported decreases in oil content in response to
preanthesis water stress, but Hall et ai. (1985) were unable to reproduce those
effects. High levels of N availability, particularly after FI or during seed filling,
may reduce seed oil concentration, although absolute amounts of oil per seed may
increase (Steer et ai., 1984). Increases in seed-protein content are often accom-
panied by proportional reductions in seed-oil content (e.g., Blanchet & Merrien,
1982; Blanchet et ai., 1983). Goffner et ai. (1988) have suggested that ABA lev-
els in seed may affect the partitioning of C between lipids and protein. The whole
issue of protein deposition in seed, its subcellular localization, its control, and the
partitioning of protein among the various functions (enzymes, oleos ins, and stor-
age protein) requires attention.
High population density increased seed-oil concentration in the four geno-
types examined by Villalobos et ai. (1994), and when seed number per capitulum
of plants grown at a single population density was reduced by limiting pollina-
tion (Steer et ai., 1988), seed oil concentration was negatively associated with rel-
ative source size (i.e., lIseed yield per capitulum). Additionally, Villalobos (per-
sonal communication) found that seed oil content showed a saturating response
over a range of seed weight generated by varying inter- and intraplant competi-
tion. In contrast, the protein and other components of seed weight increased at
least proportionately with total seed weight. One interpretation of these data is
that there is a ceiling to the absolute amount of oil that may be stored in a seed.
If availability of C during seed filling exceeds the capacity for oil deposition, then
seed-oil concentration is diluted. Rizzardi et al. (1992) have shown that cultivar
differences in the response of the hull/kernel ratio to density may be involved in
160 CONNOR & HALL
these responses. At typical commercial densities, the various effects of density on

seed-oil content may be hard to establish (Steer et aI., 1986), but they are impor-
tant aspects of seed physiology.
The preceding discussion emphasizes the many gaps in knowledge con-
cerning oil deposition in seed and the mechanisms through which various envi-
ronmental factors may operate. The temporal differences in the patterns of growth
of hull, kernel, and oil, plus the temporal spread in these patterns across positions
on the capitulum, combined with variable weighting of the contribution of the
various positions to seed yield per head, probably make an important contribution
to these effects. However, it also is clear that present understanding of the phys-
iology of the organelles in which oil, protein, and carbohydrate are stored in the
kernel must be extended if the complex interactions between the various factors
are to be understood.
Oil Type
Typically, the oil of mature sunflower seed contains approximately 110

g/kg saturated (mostly 16:0 [palmitic] and 18:0 [stearic] and 890 g/kg unsaturat-
ed (mostly 18:1 [oleic] and 18:2 [linoleic)) fatty acids (e.g., Robertson et aI.,
1978; Steer & Seiler, 1990). The proportion of oleic and linoleic acids is under
environmental, mostly temperature, and genetic control. High yields of linoleic
and of oleic acid oils are required for separate purposes in industry, so breeding
has been directed to both traits. Miller et al. (1987), Skoric (1992), and Fernandez
Martinez et al. (1989) describe recent advances in the genetic control of high
oleic acid content, Simpson et al. (1989), McCleod et al. (1990), and Downes and
Tonnet (1982) the breeding of genotypes with high linoleic acid, while Ohlrogge
et al. (1991) review the genetics of lipid synthesis in a range of oilseed species
and Mancha et al. (1994), the development of mutants with very high levels of
palmitic and stearic acids. High temperature, particularly at night, has been iden-
tified as the main environmental factor reducing the ratio of linoleic/oleic acid
content of sunflower lipids (Canvin, 1965; Keefer et aI., 1976; Harris et aI., 1978;
Silver et aI., 1984), although effects of water stress (Talha & Osman, 1975), N
(Steer & Seiler, 1990), and extremely low irradiance (Tn!molieres et aI., 1982)
also have been reported. The work of Garces et al. (1989, 1992) and Garces and
Mancha (1989, 1991) has shown that both the temperature and genetic effects are
mediated by changes in the activity of microsomal oleoyl phosphatidylcholine
desaturase. This enzyme, presumably located in the endoplasmic reticulum, is
crucial to the conversion of oleic to linoleic acid (Browse & Somerville, 1991).
Its activity is reduced by exposure to high temperature in normal cultivars and is
very low in high oleic-acid hybrids, leading to reduced oleic acid de saturation
and low linoleic-acid content of the lipids.
INTEGRATION OF PHYSIOLOGICAL RESPONSES
The discussion so far has concentrated on individual physiological respons-

es involved in the development, growth, and yield of sunflower. The elucidation
of those responses is an important first step in establishing the scientific basis of

the behavior and performance of the sunflower crop. While it is important to con-
tinue that work, it also is necessary to take a further step to investigate the inter-
actions between the component responses. This step is critical to the application
of physiological knowledge to the explanation of sunflower performance in the
field and the development of new crop ideotypes and management strategies.
Two approaches are discussed. The first concerns interactions between
pairs of responses that establish ratios and efficiencies to provide additional bases
for comparison between crops, cultivars, and management strategies. The second
concerns the development of physiologically based simulation models that seek
to provide a broad framework for the analysis of interactions between the com-
plete range of physiological responses.
Ratios and Efficiencies
Three commonly used efficiencies are the ratios of growth (or yield) to
radiation interception, water, and N uptake. In each measure, efficiency of pro-
duction can be increased by either more growth or less resource use . Efficiencies
are often as important as production, so it is necessary to know how production
is achieved. At a physiological level, it also is of great interest to learn how wide-
ly it is possible to vary linkages between what are usually closely related process-
es of growth. This provides an important perspective to the potential for the
development of new cultivars or management strategies.
Radiation-use efficiency (RUE) is the ratio of accumulated biomass to mea-
sured or estimated intercepted radiation. Two restrictions, however, should be
considered when using RUE for comparative purposes. Differences in RUE may
arise, first from differences in shoot/root partitioning and, second, from variation
in the biochemical composition of biomass. The former is a practical considera-
tion because complete root systems are rarely harvested. The latter is particular-
ly important in sunflower in which oil- and protein-rich seed make RUE intrinsi-
cally smaller after anthesis than before. For the same reason, RUE in sunflower
is smaller than in cereal species which accumulate yield principally as starch.
Two causes underlay the small RUE pa. First, a large respiratory load of the
capitulum during seed filling and second, a reduction in leaf Pn as the photosyn-
thetic apparatus degenerates during seed filling (Whitfield et aI., 1989; Trapani et
aI., 1992). The general mechanisms involved in the relationships between RUE
and leafN content have been discussed by Sinclair and Horie (1989). Two recent
studies have evaluated the effect and mechanism in sunflower. Gimenez et aI.
(1994) analyzed the nature of the response to N in the period up to anthesis. They
concluded that a fourfold response in growth to N was the product of essentially
equal effects on interception through leaf expansion and increased RUE. Hall et
al. (1995) also showed that N supply increased RUE throughout the growth cycle.
A seasonal pattern of RUE was described by Trapani et al. (1988, 1992).
They differentiated three crop stages: establishment (emergence to head visible),
rapid growth (head visible to anthesis), and postanthesis (anthesis to maximum
seed weight). Radiation-use efficiency was greatest during rapid growth (RUErg
= 2.4 g/MJ) and smaller during establishment (RUEe = 1.0 g/MJ) and after anthe-
162 CONNOR & HALL
sis (RUEpa = 1.3 g/MJ). Small RUE e also has been reported by Gimenez et al.
(1994) and small RUEpa by a number of workers (Connor et al., 1985b; Whitfield
et al., 1989; Sadras et al., 1991a; Hall et al., 1995). Although variations in parti-
tioning to roots also were involved, Trapani et al. (1992) concluded that light sat-
uration of the canopy is probably the dominant cause of the difference between
RUE e and RUE rg .
Water stress reduces RUE (Guiducci, 1988; Whitfield et al., 1989).
Nevertheless, RUE shows itself to be a conservative property in comparison with
others. In the experiment of Guiducci (1988), RUE calculated over the entire
growth period from 10 leaves to maturity was reduced by only 27% in the water-
stressed treatment (0.30 vs. 0.41 g/mol quanta for water-stressed and irrigated
crops, respectively), while PAR absorption was reduced by 61% and biomass
yield by 71 %. The difference in RUE between treatments was even smaller (21 %)
when more accurately calculated using the energy content of biomass. The con-
servative behavior of RUE can be partly explained as an effect of turgor-related
leaf movements that allow leaves to modulate the irradiance on their surfaces to
their photosynthetic capacity (Guiducci et al., 1993), as previously demonstrated
for soybean (Guiducci & Benincasa, 1994).
Intraspecific variation in RUE has been found during the postanthesis peri-
od (Trapani et al., 1988; Sadras et al., 1991a). A positive correlation between
RUEpa and seed number per unit ofpostanthesis assimilate (Sadras et al., 1991a)
suggests that sink size relative to source activity may impose a control on photo-
synthetic activity of crops with low seed set. This hypothesis is consistent with a
considerable reduction in photosynthesis of mature, slowly expanding leaves in
response to starch accumulation (Potter & Breen, 1980) and deserves further
attention.
Summarizing, sunflower leaves have high P max compared with other C3
species that does not necessarily convert into high RUE, although high rates of
crop growth have been measured in some experiments (Connor et al., 1985a;
Murata, 1981). The explanation may be complex. On the one hand, accumulated
photosynthesis over periods of days and weeks may not reflect the high instanta-
neous rates because in most environments irradiance levels required for those
high rates are relatively infrequent. On the other hand, there are many additional
steps in the progression of processes from photosynthetic assimilation to net bio-
mass accumulation. These further diminish the differences between species as
individual processes are successively combined at higher levels of spatial and
temporal organization (Gifford, 1974; Monteith, 1977; Sinclair & Horie, 1989).
For instance, RUE of sunflower was 2.2 g/MJ during the vegetative period
(Kiniry et al., 1989) vs. 1.7 g/MJ for potato (Jefferies & Mackerron, 1989). This
difference of 25% compares with a twofold difference in short-term crop Pn'
Transpiration efficiency (TE) is the ratio, most commonly of aboveground
biomass or of yield (corrected or uncorrected to production value, PV), and tran-
spiration. The physiological linkage between the two processes exists at the level
of leaf gas exchange because Pnand T occur through the same stomatal pathway.
Physiological responses that control gs or LAI (the other component of crop con-
ductance) such as irradiance, water, and N supply, affect both processes. The link-
age is strong enough to persist to the level of crop growth such that when tran-
spiration is normalized by vapor pressure deficit (Tanner & Sinclair, 1983), TE is

fairly conservative for individual species over complete growing seasons.
Transpiration efficiency of sunflower leaves calculated as the ratio of CO 2
assimilation to transpiration from short-term measurements of photosynthesis
and transpiration has been reported in a number of studies (Rawson, 1979;
Rawson et aI., 1977; Blanchet et aI., 1978, p. 12-22; Rawson & Constable, 1980).
The value of approximately 17 ng CO 2/(mg water kPalVPD) for sunflower is sim-
ilar to that of other C3 species such as wheat and cotton (Gossypium hirsutum L.)
(Rawson & Constable, 1980).
At the plant level, Briggs and Shantz (1914) included sunflower and relat-
ed Helianthus spp. in their classic study that compared TEB/T (calculated on bio-
mass) of a wide range of species grown outdoors in large containers (115 kg soil).
The values established for sunflower, categorized at that time as a "weed," ranged
from 1.3 to 1.8 gIL. Comparable values (1.7-1.9 gIL) were found for modem
hybrids grown under similar conditions over a wide range of water regimes
(Sadras et aI., 1991a). These values are comparable with other C3 species such as
wheat and cotton (1.8 gIL) and soybean (1.5 giL) (Briggs & Shantz, 1914; Shantz
& Piemeisel, 1927). Experiments using small pots (approximately 5 L) in the
field gave higher values for sunflower under limited water supply (2.3-4.3 giL)
(Blanchet et aI., 1978, p. 12-22; Green & Read, 1983).
Transpiration efficiency (calculated as TEB/T) was 33% greater before
anthesis than afterwards (Sadras et aI., 1991a). This difference arose mainly
because growth respiration of seed is greater and therefore PV is smaller than for
vegetative tissue (Sadras & Connor, 1991). When allowance was made for the
synthesis cost of oil, the difference was reduced to 13%. Most of the remaining
variation was explained by lower VPD before anthesis.
Intraspecific variation in TE has been reported at the plant (Virgona et aI.,
1990) and crop levels (Sadras et aI., 1991a). Transpiration efficiency measured
over a 32-d period for plants growing in small pots in a glasshouse, was nega-
tively correlated with C-isotope discrimination in a range from 1.5 to 2.4 mmol
C/mmol water (Virgona et aI., 1990). Seasonal TEB/T was 2.47 and 1.89 glm2 mm
for crops of a semidwarf and standard-height cultivars, respectively (Sadras et aI.,
199Ia). Seasonal differences were established in the postanthesis period and were
associated with greater RUE of the standard-height cultivar.
Nitrogen-use efficiency (NUE) can, as with the previous ratios, be calcu-
lated with either growth or yield, in this case, relative to N applied or absorbed
by plants or crops. Plant physiological interest centers on the latter while the for-
mer is more important in agronomy. It is only recently that attention has turned
to issues of the N relationships in sunflower. As previously discussed, a number
of papers have studied the effect ofN on growth, leaf expansion, TE, and partic-
ularly on RUE, but none have approached the issue from the question ofNUE.
Role and Status of Simulation Models
Crop simulation models are mathematical representations of the physiolog-

ical responses of crop development, growth, and yield to attributes of cultivars,
environmental conditions, and management. These models seek to explain crop
164 CONNOR & HALL
performance in terms of current knowledge and clearly stated assumptions about

the component processes and their interactions (Loomis et aI., 1979). In this way,
they are valuable in the synthesis of research results and at the same time in guid-
ing research by identifying areas of ignorance. The "current model-experiment-
improved model" loop is an increasingly important component of crop research.
The objectives and complexity of models vary considerably. This account focus-
es on physiologically based models designed for agronomic applications. They
have particular application in the evaluation of cultivar-management interactions
under the variable environmental conditions (e.g., Meinke et aI., 1993b; Fereres
et aI., 1993) that characterize many sunflower production areas and also can be
PTU
LAI
p/len06tage
T ratio
LAI
phenostage stem
Fig. 4-12. A relational diagram of a crop simulation model comprising three submodels. In the phe-
nology submodel, the rate of development (DVR) depends upon daylength and temperature (temp)
according to cultivar characteristics specified in terms of thermal (TU) and photothermal (PTU)
requirements for the completion of the component phenostages. In the biomass submodel, growth
depends upon temperature, leaf area index (LAI), solar radiation (solrad) and transpiration ratio (T
ratio). The partitioning of growth to the component organs (leaf, stem, dead leaf and seed) varies
during the crop cycle as specified by phenostage. In the water submodel, water enters the root zone
from where it is lost by evaporation from the exposed surface soil, by transpiration through the
crop, and by infiltration into the subsoiL The water-holding capacities of the two layers are WHCl
and WHC2, respectively. Potential evapotranspiration (pot ET) is partitioned into potential tran-
spiration (pot T) and potential evaporation (pot E). The partitioning between actual transpiration
and evaporation depends upon LAI, pot T, pot E, and the water contents of surface soil and root-
zone. The ratio of actual to potential transpiration (T ratio) specifies the effect of water supply on
crop growth. Although a simple example, this diagrammatic presentation explains the interaction
of processes in crop growth and shows how more complex interrelationships can be represented for
study and application in research and management.
used to identify targets for breeding (e.g., Sadras & Villalobos, 1994). Examples
of such applications for a range of crops is presented by Muchow and Bellamy
(1991).
In their formal presentation (Forrester, 1961), crop simulation models com-
prise sets of state variables grouped into submodels (Fig. 4-12). The state vari-
ables within each submodel describe the changing spatial and temporal distribu-
tion of a conservative quantity within the boundaries of the soil-crop system. For
example, one submodel might deal with water in various soil layers, another with
biomass in component organs, and a third with the various forms ofN in soil and
crop. An additional important submodel is that of development. This sets a phe-
nological, as distinct from temporal, timescale that identifies important events in
reproductive development such as E, FI, (BV), FA, LA, and PM. These events
signal important changes in activity of organs and hence the partitioning of bio-
mass that culminates in the expression of yield. Models use mathematical expres-
sions to represent the component physiological processes and the feed-forward
and feedback controls and nonlinear responses that characterize biological sys-
tems. For example, changing leaf growth in the biomass submodel supplies infor-
mation that affects transpiration in the water submodel. Calculation of available
soil water in turn controls both leaf growth and expansion. Nitrogen transforma-
tions in the soil determine available nitrate that adds further responses to leaf
growth and hence transpiration. The interrelationships quickly become numerous
and complex. Adherence to a formal structure in model building not only helps
to ensure internal consistency, but also facilitates model development and evalu-
ation.
Several simulation models have been developed for sunflower during the
last two decades (Horie, 1977; Anderson et aI., 1978; Chapman et aI., 1993a;
Steer et ai., 1993; Villalobos et aI., 1996). In addition, two closely related gener-
ic crop models have been adapted for sunflower (Kiniry et aI., 1992). These mod-
els differ in objectives, structure, and the number of crop and environmental
processes considered. Two models that consider comprehensive ranges of crop
processes are QSUN (Chapman et aI., 1993a) and OILCROP-SUN (Villalobos et
ai., 1996). Both consider the effects of temperature, radiation, soil characteristics,
cuitivar, and water supply on crop development, growth, and yield. The latter also
deals with N in the soil- crop system. The following brief description of OIL-
CROP-SUN illustrates the nature and application of crop simulation models to
the study of crop physiology and the application of that knowledge to crop
improvement.
The OILCROP-SUN was constructed as a physiologically based model of
the sunflower crop to investigate the comparative response of cultivars to envi-
ronment and management. It comprises virtual (i.e., not formalized) submodels
of phenology, water, biomass, and N. Information required to run the model con-
sists of daily weather (radiation, temperature, and precipitation), soil physical
characteristics, initial water and N contents, and cultivar properties defining phe-
nological development and potential yield. Using a daily time-step, the model
estimates crop leaf area, total and organ biomass, water use, and N uptake. Daily
increments in new biomass are derived from radiation interception and RUE that
responds to variations in chemical composition of the biomass, temperature, and
166 CONNOR & HALL
water- and N-imposed limitations to growth. Canopy dynamics embody the

major effects of the environment and crop population density upon leaf appear-
ance, expansion, and senescence. Actual daily growth of each organ depends
upon its temperature, crop growth rate, partitioning priorities, and water and N
limitations. Daily uptake of water and N by the crop reflect current soil contents,
root uptake capacity, and current demand. Supply/demand ratios for each provide
indices of availability that may limit crop and organ growth. The phenology sub-
model plays a central role, controlling the timing of leaf initiation, appearance,
and duration of expansion as well as floral initiation, anthesis, and the start and
150
...
G
130
:!.
110
2
U
5
"
"
90
~
G
2
5 70
Observed duration (days)

7
(b)
6
0, 5
.r::.
~
"tl
a;
4 •
'>' Sungro 380 (irrigated)
"tl 3
C Sungro 380 (rainfad)
~
:;
• 894 (irrigated)
2 o 894 (rainfad)
E
iii • Florasol
+ Contiflor3
b. Dokall G· tOO
2 3 4 5 6 7
Observed yield (t ha- 1)
Fig. 4-13. Relationships between simulated and observed values for two output variables of the sim-
ulation model OILCROP-SUN (Villalobos et aI., 1996). (a) Duration of emergence to first anthe-
sis for crops of three genotypes sown at different times over a period of 5 mo in each of two sepa-
rate years at Cordoba, Spain. (b) Grain yield of crops of SunGro 380 and 894 grown at Cordoba
under irrigation (full symbols) or rainfed (empty symbols). Also shown are results for crops of
Florasol grown over a range of plant population densities at Cordoba and of 'Contiflor 3' and
'Dekalb G-100' grown at Rafaela, Manfredi, and Parana, Argentina, during 1992 [Spanish data
taken from Villalobos et al. (1994); Argentine data courtesy of Magrin, Chimenti, Dardanelli,
Rojas, and Villar (personal communication)].
end of seed growth. Importantly, it also serves to define the timing of variations
in the priorities in biomass partitioning within the crop and, ultimately, the
expression of yield. In its present form, the model is able to mimic with good
approximation many observed features of sunflower responses, including cultivar
differences over a range of day length, temperature, solar radiation, conditions of
water and N supply, and crop population density. Some comparisons are present-
ed in Fig. 4-13.
Despite obvious limitations, the present crop simulation models are an
important beginning to an avenue of work that offers great promise to crop sci-
ence. Indeed, there is no other known way to study the interactions of processes
and provide a functionally based explanation of crop performance. For these rea-
sons, the development and testing of models will play an increasingly important
role in improving our knowledge of sunflower physiology. To the present, for
example, the development of OIL CROP-SUN has served to pinpoint many of the
gaps in our quantitative understanding of sunflower phenology across all culti-
vars for the full cycle (discussed above). It also has contributed to inspiring recent
research on leaf N dynamics and its relationship to leaf photosynthesis and crop
RUE (Connor et al., 1993; Sadras et al., 1993a; Gimenez et al., 1994; Hall et al.,
1995; Connor et al., 1995), and on the control of crop biomass partitioning
(Villalobos et al., 1992, 1994; Trapani et al., 1994). Because model building
requires explicit statements of the component assumptions, comparisons between
approaches adopted by various workers is another useful way to identify research
that may improve understanding of crop function. For example, a comparison of
QSUN and OILCROP-SUN shows distinct approaches to the modeling of root
growth and extraction of soil water, a divergence that underlines the need for fur-
ther work to quantify water extraction on general principles rather than empirical
observations of water-extraction patterns (Meinke et al., 1993b).
SUMMARY
Sunflower has long received scientific attention because its form and func-
tion have appealed to experimenters of floral development, plant nutrition, pho-
tosynthesis, water relations, and growth. It was for many years used as a labora-
tory standard to measure "wilting point" of soils and it remains an important
species in research in B nutrition of plants. However, it has been the recent devel-
opment of the species as a crop, first as open-pollinated cultivars, and more
recently as hybrids, that has been the impetus for much scientific research,
including the physiology of development, growth, and formation of yield.
Although suited to a range of environments, the crop has found a particular niche
in semiarid environments and for that reason an important additional focus in
physiological research has been in the area of stress tolerance, particularly to
water and temperature.
The first edition of this monograph did not include a chapter on sunflower
physiology, probably because little information was then available. Now there is
considerable understanding of the form and functioning of sunflower and this
chapter has attempted a structured presentation intended to be useful to those
168 CONNOR & HALL
working in all aspects of sunflower production, management, and research.

However, despite the great quantity of information available, there are still many
gaps in our knowledge of the functioning of sunflower and its response to envi-
ronment, especially under stress conditions. This review also has identified such
gaps and where possible has proposed suggestions for research, often based on
comparisons drawn with other crop species. Comparative physiology is an impor-
tant tool in research, seen in many places in this chapter, but perhaps most exten-
sively in discussions of metabolic pathways of oil synthesis. Future work in this
area is critical to success in the application of new techniques in biotechnology
to increasing crop yield and quality.
Plants are complex organisms so in consequence there are many physio-
logical processes to be considered in attempts to apply the vast knowledge that is
being gained to plant improvement and crop management. There is a real danger
that the volume of detail will obscure the parts that can now offer solutions to par-
ticular problems. On the other hand as simple problems are solved, those that
remain will require considerations of many interacting responses for their solu-
tion. Integrative models offer a means to evaluate physiological information and
to apply it to solve problems in plant improvement and crop management. The
review has shown how such techniques are now being applied with success in
sunflower research.
Much of the information presented here is new and the field is rapidly
expanding. We hope that this treatment might remain useful for perhaps a decade,
but feel confident that during that time there will be new discoveries of the phys-
iological responses of sunflower. Hopefully, during the same time there will be
even more significant development in the nature and applicability of models so
that the increasing store of physiological information can be effectively mobi-
lized and applied to solve problems and meet new challenges in sunflower pro-
duction.
ACKNOWLEDGMENTS
Our work on sunflower has been generously supported by The Universities

of Melbourne and Buenos Aires, the Australian Grains Research and
Development Corporation, and the Consejo Nacional de Investigaciones
Cientificas y Tecnicas de Argentina. These sources of assistance are gratefully
acknowledged. We thank our colleagues, Dr. Paolo Benincasa, Mr. Claudio
Chimenti, Professor Marcello Guiducci, Dr. Luis Hernandez, Dr. Peter Hocking,
Dr. Francisco Orgaz, Dr. Jairo Palta, Dr. Victor Sadras, Professor Rodolfo
Sanchez, Ms. Nora Trapani, Dr. Neil Turner, and Dr. Francisco Villalobos who
offered valuable comments on the manuscript.
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Sunflower Physiology

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Published 1997

Physiology is concerned with the development, growth, and yield of plants in

ing, which responds markedly to date of sowing and interannual variations in

Seed Dormancy and Aging

Sunflower embryos develop the capacity for germination about 6 dafter

appears to be involved in the induction of both desiccation tolerance and dor-

This phase includes two distinct subphases, germination (start of imbibition

tation of results is often difficult. Early stages in germination (radicle elongation)

Emergence to Floral Initiation

The duration ofE to FI comprised approximately 20% of the growth cycle

development by exposing seedlings to short chilling pulses (Marc & Palmer,

mated as 13 and 15 h (Hammer et aI., 1982); in another, all photoperiods includ-

Floral Initiation to First Anthesis

The duration ofFI to FA comprised approximately 38% of the growth cycle

First to Last Anthesis

Flowering in sunflower commences at the periphery of the head and pro-

Last Anthesis to Physiological Maturity

The duration of LA to PM occupied about 28% of the crop cycle of

The Terrestrial Environment

Collection of Water by Root Systems

1.8 Ra lnfed IT6) Weekly irrigated IT 1)

Plant Water Status

With the beginning of each diurnal cycle at dawn, transpiration rapidly

Understanding how sunflower responds to water shortage relative to other

Days after sowi n9

Sunflower is considered well adapted to drought, although systematic

The uptake and distribution of the three major macronutrients, N, P, and K,

120 '~-' ...

Days after sowing

Leaf area (cm 2)

Leaf area (cm 2 )

Biomass accumulates in C compounds achieved as the balance between

Leaf and Canopy Area

expression 1= Io.exp(-kL) (FigA-6). The extinction coefficient, k, describes the

0.0 +--__ -r-~-r_~-__._-~___l

Leaf area index

Seasonally, k decreases with ontogeny until about anthesis and then

Photosynthesis and Respiration

Pn = (Pmax + R){ 1 - exp[-eI/(Pmax + R)]} - R [1]

where P max is the maximum rate of photosynthesis at saturating irradiance (/), £

Specific leaf nitrogen (g N m-2)

Specific leaf nitrogen (g N m- 2)

Fig. 4-7. Relationships between parame- (e)

N concentration (SLN) (Coffi1or et a!., () °

in large pots outside in the summer in '"

between the N treatments of mean rn~

value 0.05 (SE = 0.001). (b) Maximum

Effects of Water Deficit and Nitrogen

Stems and Heads

200 400 600 800

Whitfield et aI., 1989). The exceptional photosynthetic ability of sunflower

Optimum Nitrogen Distribution

Specific leaf nitrogen (g N m-2)

Specific leaf nitrogen (g N m- 2)

Fig. 4-9. Profiles of observed (open sym- 25

tent (TNC) that would maximize crop 0

for maximum productivity will increasingly depend upon the development of

Assimilate formed in photosynthesis is the major component of plant bio-

Days after sowing

Horie (1977) pioneered attempts to describe assimilate partitioning, in

Stem 0.36 ± 0.02 0.60 ± 0.03 0.60 ± 0.03

YIELD FORMATION AND REALIZATION

crops at extremely low densities in very short-season cultivars (Villalobos et ai.,

Days after first anthesis