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Sugarcane growth and development

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 Sugarcane growth and development
Agricultural Consultant jhcock@ uniweb.net.co
Summary
This paper reviews crop management prac-
tices and changes in plant type that may b€
used to increase cane and sugar yields. Th€
lack of critical periods in the development
of the crop makes it relatively tolerant of
stress conditions. Potential yield of
improved cultivars can only be attained ifit
is matched by skilful agronomic manage-
ment.
Introduction
Much of the research on sugarcane as a crop
is now carried out by entities financed by
conmercial cane growers. Their interest in
plant physiology, and particularly in growrh
and developmenr, is principally directed to
acquiring knowledge that can contribute 10
improved varieties and malagement of the
crop. Recently Duvick and Cassman (1999)
referring to maize in the United States
lamented tlre scarcity of funds ibr research on
understanding of physiology and furthermore
noted that the limited amount is directed
mostly toward molecular approaches to
increase productivity. They went on to
observe thal molecular approaches without a
deeper understanding of physiological deler-
minants of yield potential that seek cmpirical
ly to concentrate "yicld genes are likcly 10
fail. Although Duvick's and Cassman's com-
ments refer to maize. they are dircctly appil,
cable to the sugarcane situation al prcsent.
Thcre is little iunding for physiological
research on sugar cane: the small amounts
dcdicatcd to physiology are mainly at rhe cel
lular and molecular levels with virtually no
research, outside of Australia, on growth and
developmenr ar rhe r|op ard plant lerel
Past expe ence in various crops indicales
that an understanding of the physiology of
growth and development can indeed contribuG
to increased yield. One of the most striking
examplcs is tbe ideotype for tropical rice lbr-
mulated at the Iitemational Rice Research
Institute (IRRI). This was based on an under-
standing of the growth and development of
SUGAB CANE INTERNAIIONAL, AUGUST 2OO1
by James H. Cock
tropical rice and led to the development of the
high yielding tropical rice varieties starting
with IR8 (Chandler 1969, Yoshida 1981). As a
result of this work the yield obtainable at the
plot level rapidly rose from a maximum in the
dry season of about 5 t ha' to 9 t ha,. The
results rapidly spread to farmers'fields and the
green revolution in rice began, with large
increases in rice yield in farmers'fields in
many tropical countdes.
The modifications in plant type, obrained
by breeding, were not the whole story: when
the new plant types such as IRS were com-
pared with Peta, a haditional variety, at tradi-
tional low nitrogen levels and wide spacing
the modem type showed little or no advantage
over the traditional varieries (Chandler 1969,
Yoshida l98l). Thus in the case ofrice the use
of an improved variety had to be accompanied
by changes in the management practices: This
appears to be a fairly general phenomenon
also encountered, in
maize (Duvick and
Cassman 1999, Evans and Fischer 1999),
wheat (Austin 1999), and soybe.ms (Specht et
al 1999\.
As a result of genetic improvement and
management practice avcrage farm yields
have increased in many crops over the last
iriry lcars fhe a\crage yield: ol maj/e
(Duvick and Cassman 1999) and soybeans
(Specht e/ al 1999) jn rhe USA and wheat in
lhe United kingdom (Ausrin 1999) and sugar
beet in Westem Europe (FAO 2000) have
increascd lincarly and do not appear to bc
reaching a plateau. ln terms of sugar produc-
tion. Colombia increased its productivity from
approximatcly 5 t sugar ha' yr, at the end of
the dccade of the fifties to 8 t sugar ha, y.i at
the end ofthe seventies and reached l2 t sugar
ha' yr at the end o[ the twentieth century
(Cock 2000). Furthermore there is no reason
to suspect that we have reached the limit: new
varieties with appropriate management prac-
tices are aheady producing close to 14 t sugar
hal yr' on a commercial scale on several
lhousand hectares. This situation. contrasts
with many countries where therc has been lit-
tle changc in productivity wirh world yields of
care increasing by about 207o over the last
fony years (FAO 2000). The average figures
mask large diff€rences in tendencies between
counlries. Cane production per hectare in
Australia dudng the period 1960 to the early
1990s oscillated generally between 75 and 85
t ha' in yeajs without serious weather prob-
lems, but increased to around 95 t hat in the
final years of the century (FAO 2000). For
several years Australia, which is amongst the
mosl producttve countries in terms of sugal
per ha per year was exhemely concemed with
a phenomenon they denoted as yield decline
(CapWIl et al 1991, Anon 1991) and rhe facr
that sugar yields stagnated over a period of
more than 20 years (Garside et al 799'7).
South African cane yields have stagnated over
the last forty year with no signs of al increase
in the last decade. On the other hald, ctrre
production in worlds largest cane producers,
Brazil and India, has increased steadily from
about 45 r cane ha, in the early 60s to 65-70 t
haj in the last years of the twentieth century
(FAO 2000). Nevefiheless, thc yield increases
in sugarcane have tended to be less than those
obtained in sugarbeet which has been the sub-
ject of intense and weil funded research over
the past fifty yea$: whereas world average
sugarcane yields have increased by around
twenty percent over the last forty years, sugar
beet yields have increased by nearly thJee
times as much (estimared from FAO 2000)
Whilst average yields have increased
markedly in most crops the picture is not so
clear when it comes to yield potential of rnany
crops. Evans (1993) defined yield potenrial as
the l ield of a cultivar when grown in envrron-
ments to uhich il t. adapred; wilh nulrienrs
and water non limiting; and with pests, dis-
eases, weeds, lodging and other stresses effec-
tively conrrolled. Maximum yields obtained
on experimental stations or in crop contests
may serve as proxies for yield potential.
Maximum rice yields obtained
at the
Intemational Rice Research Institute have not
increased since the first semi dwarf tropical
rice vadeties were planted in the decade ofthe
sixlies tPeng el a/ l99qr. Simllarly maximum
yields obtained in "yield contest,, in the USA
indicate little improvement in yield potential
Sugarcane growth and development
ofcom over the last twenty five years (Duvick
and Cassman 1999) and maximum soybean
yields have only recently been reaching the
levels obtained in the decade of the 60s
(Specht et al, 1999). Furthermore, Austin
(1999) found that the yield potential advan-
tage of the newest wheat varieties developed
in the nineties over the older varieties was less
than that obtained in the decade of the eighF
ies. However, Evans and Fischer (1999) in a
recent review of yield potential conclude that
there has been a progressive increase in yield
potential since the g/een rcvolution ir lhe sla-
ple crops. Irvine tlg83) reviewed the maxi
mum yields ol sugarclne obtarned in var iou"
different environments. A rapid comparison of
those yields with the highest yields obtained
more recenlly rndrcares litlle if any incredsc in
yield potential over the past twenly years. For
example in Austmlia there is considerable
excitement about the very high yields
obtained using closer row spacings (Bull and
Bull 2000), but the marimum yields obtained
in these trials are similar to those reported by
lrvine (1983) for Australia.
It appears to be a moot point as to whether
yield potential has really increased in many
crops over the last few decades, however most
crops are grown under less than perfect condi-
tions and are subjected to a sedes of stresses.
Tollenaar ard Wu (1999) suggest that com
yields in the USA have increased due to the
ability of the new varieties to capture
resources more efficiently, and that these dif-
ferences only manifest themselves under
sftess. Average yields of many other crops
have increased substantially, at the farm level,
over lhe past few decades and lhere is no e\j
dence to indicate that farm level yields are
reaching a plateau and may not continue to
increase in the loreseeable future if suitable
vadeties are produced and adequately man-
aged.
In this paper we will analyze crop man-
agement practices and changes in plant type
that may be used to increase farm yields. First
we will describe in general terms the process-
es involved in crop production of sugarcane
and other crops. We shall then describe the
growth and development of sugarcane under
"standard conditions" and how changing these
conditions affects the different growth and
development processes. From there we will
look ar the basic processes lhal determine
yield, which are radiation irterception, photo-
synthesis and partitioning of dry matte. Then
based on this we shall speculate on how to
further increase quality and productivity in the
future. During this process examples wiil bc
given from sugarcane and other crops. We
will not enter into detail at the cellular and
molecular level as these areas were consid-
ered to be outside the scope of this review and
have been covered in other fora.
Growth ond development
Crop production is essentially a very simple
sysrem driven by photosynrhesis. Loomis
(1999) indicates that the key elements in the
system are the inlerception of photosyntheti-
cally active radiation, use ofthat energy in the
reduction of carbon dioxide and other sub-
strates (photosynthesis), incorporation of the
assimilates into new plant structures (biosyn-
thesis and growth) and maintenance of the
plant as a living unit. Achieving high yield is
conceptually simple: maximize the extent and
duration ofradiation interceptionl use the cap-
rured enere) in efi:crent p\oto.)nlhesrl: pani-
tion the new assimilates in ways that provide
optimal proponions of lcaf,stem, root and
reproductive sttuctures; and maintain those at
minimum cos!. Allhough Llus scheme rs qurn-
tessentially straightforward, the details aie
extremely complex and the dynamic aspects
of partitioning particularly intractable
(Loomis and Amthorl999).
Crops can basically be divided into two
major categories in terms of their growth and
development. Sequential development crops
first develop photosynthetic capacity and the
organs that are economically useful, and then
the photosynthetic capacity is utilized to fill
the economically uselul parts. Simultaneous
development crops on the other hand develop
pholosynthetic capaciry ard lill the economi-
cally useful parts simultaneously, at least dur-
ing a considerable peiod of their growth and
development (Cock 1983, 1984, El Sharkawy
and Cock 1987, Moore 1989).
The cereal crops are typical of the sequen-
tial development crops and essentially have
t$o distinct and growth phasesr vegetative
and reproductive with the latter divided into
pre-heading and post-heading. Yield capacity
in these crops is primarily determined in the
pre-heading period whilst ultimate yield is
determined in the posrheading period
(Yoshida 1981). During the vegetative phase
leaf area builds upr following induction of
flowering and beginning of the reproductive
phases the remaining leaf primordia develop
followed by the inflorescence. By heading
leaf growth has essentially ceased and grain
growth begins. This sequential development
trait leads directly to two imponant character-
istics of these crops. Firstly. yield tends to
SUGAR CANE INTERNATIONAL, AUGUST
increase with the LAI at heading reaching a
plateau above which it remains constant
(Yoshida et al 1972). Secondly, sequential
crops often have critical periods of develop-
ment when a shofi period of stress can cause a
marked reduction in yield or if severe cause
complete crop loss without causing plant
death. Examples are stress in the reduction
division period in rice, which causes spikelet
sterility in rice (Yoshda l98l) or in the silk-
jng to anthesis period in com, which marked-
ly reduces maize yield (Bolafros and
Edmeades 1996).
Simultaneous crops such as sugar cane,
oil palm,sugar beet and cassava simultane-
ously de!elop ne$ phorosynthe c capacjl)
whilst they are filling the economically useful
duing a substantial portion of
organs, at least
their growth cycle. These crops tend not to
have critical periods when stress leads to
severe yield reduction or even crops failure.
Furthermore dunng lhe lillinB period lhere is
a balance between the formation and mainte-
nance of photosynthetic capacity and the fill-
ing of the economically useful organs. If par-
titioning towards the leaves is excessive then
little assimilate is diverted to the yield organs
and hence yield tends to be low, or if most
assimilate is utilized in the economically use-
ful parts then the photosynthetic capacity may
be reduced to such a low level that yield is
also reduced. Hence simultaneous develop-
ment crops tend to have an optimum leaf area
index at which yield of the economically use-
ful part is maximized (Cock 1983, 1984 and
El Sharkawy and Cock 1987, Corley 1983).
Many ofthese crops also have no well defined
maturity periods.
Sugdrcane i5 a relativcl) simple crop in
terms ofdevelopment. Although it may flower
and produce seed this is not necessary for
commercial production. Basically the growth
ald development of the crops consists in the
growth ofpurely vegetative organs, stems and
roots. [t has no critical periods ofgrowth such
as anthesis in the cereal crops when a short
stress can cause disasffous reductions in yield.
The sugarcare crop can be considered prima-
dly as a simultaneous crop, however, it also
has some aspects of a sequential crop. In the
early period of development final shoot num-
ber (an important determinant of yield capac-
ity) is established in the early phases and leaf
area is produced with little simultaneous accu-
mulation of sugar in the stalks. Nevertheless,
once stalk elongation begins there is simulta-
neous accumulation of sugar in the stalks and
maintenance and development of photosyn-
thetic capacity.
2OO1

Crop development
The sugar ca.ne crop is normally grown as a
perennial crop and is harvested several times
before replanting. The hrst cycle is referred to
as the plant crop and subsequent crops as
ratoons. Although sugar cane can be grown
from true seed, commercial plantings are
always made using stem cuttings or setts often
called seed.
Seed and germikation
The planting mareial is taken from elongated
stalks ofcane plants and is normally placed in
the bottom of a furrow and covered with soil.
Roots form on the intercalary region and the
axillaiy buds germinate. There is strong apical
dominance in this phase and the germinating
buds nearest to the apical end of the oiginal
cutting suppress the germination ofnew buds.
Time to sprouting from the seed piece differs
between va eties and is temperature dep€nd-
ent occurring after approximately 350.C days
using a base temperature of 9.Ci (Keating €t
al 1999).
In ratoon crops axillary buds from the
original stool germinate after about 100"C
days (Keating ?r at 1999).
Tillering
The newly sprouted plants enter into a rapid
tille.ing phase with new shoots being formed
from the a,rillary buds at the base of those
shoots that are already established. Leaf
appearance rate per shoot is relatively rapid at
this growth stage. ln the Colombian sugar,
growing region with an average temperature
about 24oC a new leaf appears approximately
once every onc to two weeks on each shoot. ln
Australia in simulation models the leaf
appeaJancc ratc at lhis earl) gro\4lh \taee is
one new leaf every 80"C days, which is faster
than in Colombia. The shoots have a high
Leaf Area Ratio (leaf area per unit total shoot
weight) at this stage. Leaves are still relative-
ly small and leaf longcvity short. oftcn being
less than one month from initial appearance to
senescence. The rapid tillering and develop-
ment of new leaves is associated wilh the
rapid uptake of nitrogen and other nutrients.
The tillering phase terminatcs when
murual \hading occurs and slems begtn to
elongatc (lrvine 1983). Many of thc smaller
tillers die and total shoot number is reduced
by as much as 507. (Bull and Glazsiou 1975).
It appears that as rhe tillers become
aulonomous. those that are shorl and shaded
SUGAR CANE INTERNATIONAL, AUGUSI 20O1
cannot provide sufficient photosynthate for
their own development die. Bull and Glazsiou
(1975) suggest that although the reduction in
tiller numbers may be large, the LAI that is
lost is insignificant in development terms.
The excess tillering and subsequent regula-
tion of the number of shools rhar survive is a
plastic response that is agronomically desir-
able. Caps in the stand will be compensaled
for as there will be less shading and hence
less tiller death.
Stalk elongation to maturity
Stalk elongation commences 1200 to 1800"C
days after emergence (Keating et al 1999).
During the period ofstalk elongation and mat-
uratron the number ol slalks normally remains
constant. In Australia there are reports ofcon-
siderable stalk death during rhis phase
(Muchow er dl 1995). Nonetheless, other
reports in Aushalia have not substantiated a
generalized phenomenon of stalk death,
although there was:m association between
stalk death and lodging (Singh er a/ 2000).
The leaf appearance rate per stalk
declines during this peiiod, whilst the leaf
size of new leaves increases and reaches a
maximum as late as nine months after plant-
ing (lrvine 1983) after which ir stays close to
the maximum or decreases slightly. In
Colombia we have found individual leaves of
850cm' in CC 83-25, bur 500-700 cm: leaf.'
is more common. In Mauritius Cheero
Ndyamarh el ,rl t2000) recorded meximum
leaf sizes of 650 cm, at this growlh stage,
with some varietics with smaller leaves ar
400 cm'1eafi. Nitrogen content declines both
on a per unit leaf area and per unit weight
basis. The leaf longevity is much greater in
this growth stage with leaves often living for
5 months after appearance (lrvine 1983).
Amaya et al (1985) report Ieaf longevities of
140 days for leaves that appear when the
plant is seven months old. The net effect of
these changes in the developmcnt pa[ern is
that leaf area per stalk also tends to decline
slowly as the crop matures. ln young plants
in the tillering (15 weeks old) phase leaf
nitrogen content may be as high as l.? gN m
: but dcclines to
0.85-l.2 gN m-, by 45 weeks
after planting and may be as low as 0.8 gN m
'?in oldcr plants (Allison er al 1997. Wood et
al 1997). The reduction in leaf appearance
rale. little change in leaf size and stalk elon-
gation is associated with changes in the par-
titioningol assimilate and deposition of
sucrose, which commenceS in the lower
nodes.
Sugarcane growth and development
The dry matter content of the stalks
increases from about lSVo when elongation
commences to about 3070. Sucrose as a per-
centage of the total dry matter increases and
reaches levels up to 5570 of the dry matter
(Bull and Glazsiou 1975). Muchow et al
(1996) indicated that the suqose as a percent-
age of the stalk dry matter was relatively con-
stant at 48% across varieties and suggested
that there was little evidence of breeding
progress in this character whilst data from
Cenicaffa (2000) indicates consistent differ-
ences between varieties in this character
The cane plant is indeterminate and does
not matue and cease to grow. Under the rela-
tively constant climatic conditions of the
Colombian sugar producing region where
average monthly temperature varies by less
than l'Cthe rate of biomass production
declines, but does not approach zero, when
the crop reaches approximately 12 months
(Cenicafla 2000). The extenr of the decline in
biomass produclion varied in lhe free \ari-
eties studied. lndividual shoots continue to
produce new leaves and gain weight, however
in some cases total biomass productiot may
reach a plateau due to death of some shoots
(Muchow et al 1995). Singh et at (2OOO\ did
not observe this type ol plateau response, but
they did note greater stem mortality when the
crop lodged.
Flowering
Flowering is generally considered as an unde-
sirable characteristic in commercial cane
fields, but is necessar-y for breeding programs.
Floraenng occurs rn some vaneties of cane
under certain conditions. principally shorten_
ing day length causes the induction of flower-
ing. For induction to occur the plant needs to
be well watered. The apex becomes reproduc_
tive eventually produces inflorescence and the
shoot will eventually die.
Ir,terceptioh and conversion of solt r
radilttion
Solar radiation is rhe driving force behind
photo\)nthesis. ! hrch dcLermtnes lhe pri.
mary producuvit) of crop slands. In lhe pen_
od immediately after the second World War
the concept of LAI, ratio ol leaf area to land
area was developcd (Watson 1952). The radi-
ation intercepted and used for photosynthesis
is primarily determined by the LAl. A LAI of
L AII information presented in this paper on degree
days is ba.ed on rhe uorl ol Kealrng ?r o/ ( lqgql
and uses a base temperature of 9"C
Sugarcane growth and develoPment

one distributed in a perfect horizontal mosaic development do not even attempt to simulate leaf growth and this is associated with a
tillering; standard data sets arc used for shoot decrease in the sucrose content of the stalks
would be capable of intercepting all the
number. Considerable work was carried out (Bull and Glazsiou 1975).
incoming radiation,'but most plants do not
on tillering in wheat, barley and rice many The total number of expanded functional
have a perfect mosaic and leaves ale not hor-
years ago: the general observations at the leaves per stem normally fluctuates some-
izonlal. Funhermore. the quantum efficiency
field level suggest that the same factors that where between 6-13 (Bull and Glazsiou 1975'
of photoslnthesis ol individual leaves is
control tillering in these crops also influence Amaya et al 1995, Cheero-Nayamath et al
greatest at lower photon flux densities and
tilledng in sugarcane. In general tillering 2000, Keating et al 1999, Cock et al 1'99'l).
hence radiation use efficielcy in a plant com-
increases with higher levels ofnitrogen' high- The leaf number per stem in a well developed
munity will be greater when a larger leaf area
er temperatures, increased solar radiation and crop is probably regulated by shading: if you
rcceives a lower photon flux density than
is restricted by water deficits. Normally walk up to a cane field, from the roadside you
when a smaller area is intensely illumilated.
plants produce many more tillers than they will see a dense wall of green leaves often
Thus not only is leaf area index important but
extending to ground level, but ifyou walk into
also the disposition of the leaves in the require, however those tillers that are shaded
the crop you will only find green leaves on the
canopy (Monsi and Saeki 1953). The con- excessively die (Cock 1969) Due to the pro-
cepts of LAI and leaf disposition have been duction of excess of tillers, stresses at the uppermost nodes.
early growth stage can often be compensated Plant crops have a slower build up of LAI
greatly refined since the early work referred
for later on. In sugarcane the number oftillers than ratoon crops, but eventually reach greater
to here. ln spite of the complexity of the
was reduced by early season water stress, but LAIS. Although the phenomenon of lower
analysis required to understand and model
the final shoot number, LAI and yield was LAI and the associated reduced growth rates
leaf canopies (see for example Anderson
similar to well watered controls (Robertson et of ratoon crops are extremely well known' no
19&t) the conclusions that can be drawn are
al 1999a). plausible and satisfactory explanations are
relatively straiShtforward Canopy photosyn-
thesis at low LAls is maximiled by horizon- The rale of leaf appearance per stem ls available.
reduced by water deficits and leaf senescence It should be noted that LAI development
tal leaf dispositions that intercept more radia-
accelerated. Funhermore the leaf expansion of cane is slow compared with many other
tion. At low LAIS increased canopy photo-
rate is exffemely sensitive to water deficit. crops under similar conditions You don't
synthesis is nore likely to be obtained
Leaf expansion is reduced at much smaller need to be a physiologist or take measure-
lhJough increased leaf area lhan changing
water deficits than those that directly effect ments to see the difference between the
leaf disposition. As LAI increases beyond
stomatal conductance and leaf photosynthesis ground cover of cane two months after plant-
about three, mole verlical leaf disposition
(Roberts et al 1990). Inman-Bamber and ing and that of maize or soybeans at the same
becomes more efficient in a C 4 crop such as
Jaeger (1986) indjcate that sugdcane com- age. Various authors have commented on the
maize (Loomis and Williams 1969) The
high- pensates with accelerated growth of new slow build up of LAI (lrvine, Benda and
advantage of ve ical leai disposition at
elevation at mid leaves when water deficit is relieved, howev- Ricard 1980, Muchow el al 1994, Bull and
er LAls is greater when solar
(for in trop- er, Robertson et al (1999^) indicated that in Bull2000).
day approaches vertical example
mid season water stress leaf size was reduced The cane leaf canopy is relatively erect
ical conditions) and when most of the radia-
and new leaves that appeared when the stress and Muchow et al (1984) indicated a light
tion is direct solar radiation: under cloudy
was removed continued io be small for a pro- extinction coefficient (k) of 0.38, which is
conditions with diffuse radiation particularly
longed period. commensurate with an erect habit. There are
at higher latitudes leaf disposition is of less
Tempemture effects on leal development dis(inct differences between genoptypes,
importance.
are nol well documenled Bull and Claz"rou however. Irvine and Benda (1980) did not
(1975) recorded little difference in total dry hnd differences in growth rate relaled to leaf
Leaf area index and ditposition of the IeaJ
matter production with tempe.atures in the disposition. ln several trials Muchow cr al
canopy
range of l5-30"C in the first six months sug- (1994) found only 6070 of total radiation
LAI development in gestlnB relatively lrllle effect on leal aJea intercepled in growth cycle. With these rela-
The general pattem of
development over this range, however as tem_ tively low levels of radiation intercepted it is
sugilrcane is to increa\e up until a maximum
peraturc increases shoot to root ratio also probable thal LAI ls lhe ovenrdrng faclor in
somewhere between 4-8 months and then to
of increases. Neve heless personal visual obser_ determining growth rates and leaf disposition
slowly decline. M&{imum values more
vations on the growth of the same varieties ai is of secondary impo ance However, if it
than 8 have been reported, but more common
differenl elevations, but similar latitudes, sug- were possible to incrcase LAI (particularly
values lie between 4-5 (lrvine 1983)' although
gest that tillering and leaf appearance rate early on) then leaf disposition would become
Keating el 4l (1999) in an analysis of a series
increase considerably as temperatue increas_ more important.
of experiments in diflerent parts of Australia
es in the range of 15 to 30'C and leaf area Lodging in most crops radically changes
repon a high proportion of the trials with
development is more rapid at higher tempem- the disposition of the leaf canopy, but there is
ma-ximum LAIS in the 6-8 range.
of leaf appearance rate' tures. Bull and Glazsiou (1975) point out that little or no quantitative information available
LAI is balance
Ieaf expansion is very slow beiow 15'C. on the effects. Visual obseNation of lodged
leaf size. and leaf senescence. Leaf appear-
Kealtng et al (1999) use a base temperature cane crops Indicate lhdt the leal disposition is
ance rates depend on stalk number or tillers
rate per tiller' for shoot development of 9"C suggesting that more planophile than in non lodged crops
and leaf appearance
Surprisingly little is known about tillering rn below this level new leaves aJe not formed
When tempentwe increases, particularly in 2. Some ofthe radiation inlercepted might be either
cane. Most of the models now being reflected or transmilted through the leaf.
developed to simulate cane growth and maturing crops, fiere may be a flush of new

SUGAR CANE INTERNATIONAL, AUGUST 2OO1


Furthermore the vertical distance between
leaves is greatly reduced.
Pholosynthesis, crop growth rate and
rad.iatio n use effc iency-
Sugar cane is a C 4 plant and photosynth€tic
rate is among the highest found in crop plants
(Irvine 1966,1975,1983). Vadous authors
repot large variations in the photosynthetic
rate of different vaieties (see Irvine 1983),
but the differences tend to be small when
plants are grown under standard well con-
trolled conditions (Bull and Glazsiou 1975).
Whilst varietal differences ate small, there arc
very large differences in photosynthetic rate
ofindividual leaves related to the age of plant
(Had and Bun 1967, Waldron et al 196'1,
Kortschak and Forbes 1968, Bull and Tovey
1974, Amaya et al 1995, Allison et al 1997)
wrth photoslnthcUc rate declinrng lrom ma\i-
mum levels ol
about 45 pmol m: s-' on
intensely illuminated young leaves on three
month old plants to about 25 ,.mol m, sr on
ten month old plants ( Amaya et al 1995). The
decline in photosynthetic rate is associated
with a marked decrease in lcaf nitrogen
(Allison el al 1997, Cenicana unpublished
data) and with reduced stomatal conductance
(,\maya et al 1995). The phorosynth€tic rates
of leaves with sirnilar nitrogen levels but from
plants of different ages are essentially similaJ
(Allison el al 1997) suggesting rhat nitrogen
con(ent is a funddmenLal laclor tn determinlng
the decline in photosynthetic rate associated
with older plants. Wood er rll (1996) indicare
that photosynthctic rate only declines $'ith
leaf nitrogen when this is below 1.2 gN m:,
the figure used by Sinclair and Horie ( 1989)
for maize as a C 4 plant. lt should be note that
in the original data presented by Sinclair and
Hode (1989) there arc no dara points above I
gN m and they also report that lhere are vir-
tually no respon5e studies belwcen leaI nrlro
gen and photosynthetic rate for maize. On the
other hand the data ofAllison ?/ 4l (1997) for
sugarcane clearly indicate that photosynthetic
rate atborh low and high pholon l'lux den\itres
with increasing specific ieaf nitro-
increases
gen content up to the maximum levels
obtained of 1.7 gN m:. Furthermore, there
was no evidence of a leveling off of the
response at 1.7 gN m, and they suggested that
crop photosynthesis is commonly restiicted
b) the lou niLropen le\cls ot rhe ledvcs during
much of the life cycle.
At the canop) le\el r!dtJLion usc cfftcren-
cy (RUE) has commonly been utilizcd to
determine ho$ effecti!ely a crop uses the
SUGAR CANE INTERNATIONAL, AUGUST 2OO1
intercepted radiation. The RUE can be defined
as the biomass produced per unit solar radia-
tion intercepted by the cropt as such it can be
used as a proxy for gross photosynthesis
minus respiration. The carbon assimilation of
a crop stand does increase as the total radia-
tion increases with no indication of light satu-
ration, however, the increase is not linear
indicarrng that there may be problems with the
utilization of RUE (Asseng and Hsiao 2000).
Loomis and Amthor (1999) question the utili-
ty of RUE whilst at the same time acknowl-
edging that altemative methods for study of
canopy photosynthesis are diificult or expen-
sive. At the same time they note that crop
modelers [who dominate the crop physiology
area in sugar cane at present] are reluctant to
use simulation models of canopy photosyn-
thesis to replace RUE.
Sinclair and Horie (1989) suggest rhat
RUE is extremely sensitive to changes in leaf
photos) nthetic rate and rhal major declines in
the photosynthetic rate resulting in stresses or
crop maturity will result in substantial losses
in crop biomass production, whilst increases
in the maximum photosynthetic mte greater
than those normally observed are unlikely to
increase RUE substantially. The RUE of sug-
arcane under favorable conditions. before bio-
mass accumulation begins to falter as the crop
ages, is estimated at 1.'72-1.'15 g MJ' in rhe
plant crop and slighdy less at 1.59 in rhe
ratoon crops (Muchorv el al 1994, 1991,
Robefison e/ d/ 1996. Wood er a/ 1996). After
the boom pcriod of growth RUE falls marked-
lyand it has been suggested lhat this decline
and the lower RUE associated with ratoon
crops is not related to the lower nitrogen con-
tent of the leaves (Muchow e/ a1 1994. Wood
€/a/ 1996). Nonetheless, we hale noted above
that declines in photosynthetic mte related to
stress will
substantially reduce crop biomass
production (Sinclair and Horie 1989) and also
that leaf photosynthesis declines with declin-
ing Ieaf nitrogen content associated with crop
age. Hence, it seems most probable that the
severe reductions in RUE with crop age and
the lolver RUE in ratoon crops are directly
related to the lower nitrogen content and
hence photosynthetic capacity of the leaves
thal form the crop caropy. lt also appears like-
ly that the lower RUE ofmtoon crops is relat-
ed 10 the lowcr nitrogen content of the leaves
normally found in ratoon crops.
Water stress at the field level dramatically
decreases RUE (Robe son et al 1999a\. The
effect is presumably mediated by a reduction
in the photosynthetic rate of the individual
leaves in thc canopy rclaled to decreased
Sugarcane growth and fuvelopment
stomatal conductance. The mechanisms that
control stomatal rcsponse to water stress are
complex, involving direct effects of water
potential on the stomata, root signaling and
leaf to air vapor pressure deficits inter alia.
The most commonly recognized control
of stomata by water stress is a direct response
to the water status of the leaf. When leaf water
potential falls below 1.3-1.4 MPa the stomatal
conductance begins to decrease, reaching
extremely low levels that rcsult in negligible
photosynthetic activify at 1.7 MPa (Roberts e,
al. 1990).
The stomata of many troprcal crop species
respond directly to the leaf to air vapor pres-
sure deficit (VPD) (El Sharkawy €r al 1984).
Sugarcane is no exception: the leaf conduc-
tance is reduced by 25-5070 of that at a leaf to
aii VPD of 0.5 kPa when leaf to air VpD
increases to I kPa. which is a level that can be
encountered under field conditions (Grantz
and Meinzer 1990, 1991). It is possibie that
the direct response to leaf to air VPD in sug-
arcane may in some circumstances limit pho-
tosynthesis and hence biomass production.
Increased crop yield has been obtained in
well-watered cassava, a highly sensitive
species, by increasing the air humidity and
hence reducing leaf to air VPD (Cock et al.
1985). whilst sensirivity to leaf to air VpD
has been suggested as a mechanism that is
advantageous under conditions of water
delrcrt ard ma1 lncrea\e water use efficienc)
of a crop, under well-watered irigated condi-
tions sensitive stomata will reduce crop pro-
ductjvity.
Leaf photosynthesis of wheat may
respond to water stress in the root zone even
when there is no measurable change in leaf
waler potential, due to root signaling mediat-
ed by plant hormones produced in the roots
(Ali er al 1999). Meinzer er al (1991) provide
.onvrncing e\ idence lhal similar mechanisms
exisl in sugarcane and that stomatal conduc-
tance is modulated by root signals.
Furthermore, Meinzer and Crantz (1990)
manipulated leaf area per plant and demon-
strated that stomatal conductance decreases
wjth leaf area per plant in such a manner that
there is an asymptotjc relation between total
canopy transpiration and LA[.
The C 4 plants tend to be water use effi-
cient, and sugar cane is no exception produc-
ing about 7-9 kg biomass kg water, (Bull and
Glazsiou 1975). Based on a haryest index of
0.39 (Muchow er al. 1996), 1000 rnm of rran-
rpirarion r10.000 r waler ha r should give a
biomass yield of 70-90 t ha' and a sucrose
yield of 27-35 t sucrose ha,. ln the Cauca
Sugarcane growth and development
valley in those areas where crop transpiration
is estimated to be of the order of l0O0 mm,
sucrose yields in variety trials aje frequently
in this range (Cenicaffa 1997). Simple analy-
sis of this sort should be used with caution:
water use efficiency (WUE) declines linearly
with increased VPD and hence comparisons
between areas and absolute assessment of
WUE should include a reference to the VPD
or some proxy such as the potential evapo-
ftanspiration, however even then interpreta-
tion is not easy (Asseng and Tsiao 2000) and
unlortunately the required information is nor-
mally lacking.
Maximum recorded crops growth rates of
sugar cane arc in accordance with levels
expected for C4 crops. Muchow dl. (1994)
"r
recorded 41g m' day' for a period of 20
weeks and various authors report similar lev-
els over shorler trme penods. These malimum
growth rates occur when LAI is sufficiently
large to intercepl most of the incoming radia-
tion (eg 907o or more) and before growth rates
decline due to the crop getting old.
Part ioning
In the initial tillering phase of growth the
assimilate is partitioned between the roots and
the shoots, with the shoot growth beinS parti
tionedhetween new tillers. and leaves and leaf
sheath with a minimal proportion passing to
the stalk. ln this phase as much as 3070 of the
biomass goes to the roots (Keating et al 1999).
water stress and long photoperiod both tend to
increase the root to shoot ratio (Bull and
Glaz siou 1975).
As noted above the tillering phase termi-
nates when mutual shading occurs and the
stems begin to elongate (lwine 1983). The
stalk elongation begins when total above
ground crop biomass is around lOt ha1
(Robe son 1999a). During this stage approx-
imately 707, of the new biomass accumulates
in the elongating stalks, and most of the
remainder in new leaves and the top part of
the plant denominated "cabbage" by the
Australians (Keating er al 1999). At about 300
days after planting 60-'/0Vo of the standing
aerial biomass (not including fallen leaves)
was found in stalks increasing to close to 907,
by 450 days after planting (Cenicana 2000).
During the stalk elongation phase the
sucrose percentage dry matter increases and
reaches a plateau level with up to 98% of the
total sucrose in the stalk (Cenicafla 2000,
Muchow et a, 1996). The maximum stalk
sucrose percentage dry matter rn a comparison
of three vaiieties was 5570 in the high sucrose
10
vaiety and 41Vo in the average sucrose van-
ety (Cenica.fia 2000). The stalk sucrose dry
matter percentage for a given variety appeajs
to be relatively stable and vades little under
stress conditions. When planls were severely
defoliated the stalk sucrose dry matter per-
centage was not affected (Cock et al 199'7).
Similarly water shess in the pe od before har-
vest has relatively small effects on stalk
sucrose dry matter percentage (lnman-
Bamber and Jaeger 1986, Ellis and Lankford
1990. Robertson and Donaldson 1998,
Robertson, Muchow and Wood 1999), howev-
er even small differences can be commercial-
ly useful.
Although partitioning in crop plants nor-
mally refers to partrlroning of dry biomass. in
the case of sugarcane, where transport of the
fresh product is a major item in the total cost
of the final product, the partitioning between
dry matter and water in the stalk is critical.
The dry matter content of stalks may be as
low as 107, (Keating ?l al 1999) when they
begin to eloogate, but by maturity 3070 dry
matter is a fairly standard figue with about
half the dry matter as sucrose. Drying off
before harvest increases the dry matter con-
tent of stalks (Robertson, Muchow and Wood
1999). As in the case of sialk sucrose dry mat-
ter percentage the effect is quite small, never-
theless it may be of commercial significance.
During the final stages of growth before
haryest the sugarcane crop acts as a simuha-
neous development crop in which maxlmum
production is achieved by a delicate balance
between partitioning assimilate to leaves so as
ro ensure asslm)lale rupply and nol divening
so much assimilate to the leaves that there is
lirrle left to fill the stalk with sucrose.
Increases in temperature that lead to new leaf
growth (Bull and Glazsiou 1975), new flushes
of leaves when water stress is alleviated
(Robertson, Muchow, and Wood 1999) or
growth of axillary buds on lodged cane may
alter partitioning in favor of new leaf devel-
opment and reduce yields. Drying off and
treating cane with ripeners such as glyphosate
reduce the growth of ieaves :Lnd may not only
increase the sucrose concentration but also the
overall sugar yield per hectare (Lairahondo
and Villegas 1995).
Opportunilies to improve the cane crop
In a potential productivity symposium in S.E.
Asia in 1980 Irvine (1983) commented, 'The
surest prediction for sugarcane is that yields
will continue to rise if breeders, production
agronomists, plant protection specialists, and
engineers continue to contdbute new informa-
tion to crop production managers." I f€el that
statement is as {rue today as it was twenty
years ago. Let us now speculate, from a phys-
iological standpoint, on some of the means
that the cane crop can be improved in the
coming years.
The improvement of the cane crop is not
as simple as improving the total yield of
sucrose per ha. The crop has to fit into man-
agement systems that may be restricted by
technical, social and environmental con-
straints (Cock et dl. 1999); the plofitability of
different production systems is extremely sen-
sitive to the sucrose concentntion in the har-
vesbd product (Cock et al 2000); and the
chanctenstics of the varieties grown must
facilitate such costly operations as harvesting
(Cock et al 1993). Firstly the question of
increased yie)d is addressed and then other
desirable characteristics such as sucrose con-
tent and ease of harvest.
IncreasinB yield.
There are basically two ways to increase yield
of a crop: increase total biomass or incrcase
the economically useful part of fte total
biomass.
lncreasing biomass production
lf we accept that respiration is a necessary
part of the growth process and that it will be
difficult to reduce respiration as a proportion
of total photosynthate produced, then total
biomass itself can also be increased essential-
ly by two means: increased interception of
solar radiation or more efficient use of that
solar radiation in phofosynthesis.
Radiation interception
As we have noted previously, the LAI of the
sugarcane develops relatively slowly and
there may be opportunities to increase yield
via increased interception of solar radiaiion
(lrvine et al 1980, lrvine and Benda 1980,
Keating er al 1996, Bull ard Bull 2000a,
2000b). Two of the most obvious options for
increasing LAI and hence radiation intercep-
rion are planting at higher densities and
increased nitrogen applications. At present in
Australia there is considerable enthusiasm
(one might even say furo.) for the route of
high densities to increase yield (Ridge and
Humey 1994, Bull and Bull 2000a, 2000b).
Nitrogen has also been used to increase yield
in cane, however a point is often reached
SUGAR CANE INTERNATIONAL, AUGUST 2OO1

where increased nirogen leads to an undesir-
able reduction in sucrcse concentmtion
(Larrahondo and Villegas 1995) of the cane
and lodging. In various other crops higher
densities and nitrogen applications are part of
the package that has lead to higher yields
(Chaxller 1969, Duvick and Cassman 1999,
Evans and Fischer 1999). Nevenheless, in
order to obtain higher yields at higher densi-
ties and nitrogen level, varieties with specific
characteristics were required.
Lodging is associated with high LAIs and
biomass production. In the case of the cereal
crops lodging resistance is essential
if yield is
to be maintained al high densilies and nitro-
gen levels as when lodging occurs biomass
production is adversely affected (Chandler
1969, Yoshida et al19'72, Rajcan and
Tollenaar 1999, Austin 1999). Similarly an
important characteristic of new sunflower
varieties is lodging resistance (Lopez Percira
et al 2C0O). In sugarcare Singh sr al (1999,
2000) have recently demonstrated that when
Iodging occurs crop growth decrcases. In sug-
arcane lodging is associated wirh high bio-
mass as heavy cane tends to lodge, whilst at
the same time lodging is a constraint
obtaining even higher biomass. Nevefiheless
there are large vadetal differences in lodging
aesistance
Although increased nitrogen fertilizer
increases leafarea it may reduce sucrose con-
tent of stalks. Applying potassium amelio-
rales this effect in some varjettes tQuintcro
and Garcia 1997). Amaya (pers. comm.) grew
a large number of varieties at high and low
nitrogen levels and found that some \ ariet.es
mantaned lugh sucrose conrenl at high nilro-
gen levels indicating that it may be possible
to select varieties that tolerate high nitrogen
levels.
P hoto synt hetic eff c ie ncy
EfforLs Lo improvg crop yrelds b1 increasing
the ma\imum photosynthetic efficiency have
generally not been successful. Loomis and
Amthor (1999) note rhar dudng evolution
photosynthetic systems have been higlrly con-
served and domestication and crop improve,
ment ha\e not increased rhe basic efficiency
ofthe process. Nevefiheless there do appear to
be opportunities to increase the efficiency of
photosynthesis of crop canopies; particularly
in those periods when at present they function
at low levels of elficiency.
The nitrogen content and the photosyn-
thetic eft-icienc) ofcane leaves declines rapid-
l) as rhe crop ases and this is reflected in
SUGAR CANE INTEBNATIONAL, AUGUST
reduced radiation use efficiency (RUE). One
of the major factors involved in increasing
yields of several crops is the stay green char-
acter in which leaves do not senesce rapidly
when the crop matures (Evam and Fischer
1999). This character is important (Duvick
and Cassman 1999) increasing total biomass
production in maize (Rajcan and Tollenaar
I9qga. lggqbr and soybeans (Specht ?r ;1
1999). Fischer at al (1998) indicate that more
efficient photosynthesis in the seed filling
period is associated with greater biomass pro-
duction and yield in wheat.
I suggest that a concentrated effort to
seiecl "slay green cane larieties that main-
tain their nitrogen levels as the crop matrres
offers an interesLing posstbiliry for increasing
photosynthetic efficiency of sugarcane and
increasing yield.
Tn a senes of expenments at Cenicatia
almost all the variation in photosynthetic rate
of individual leaves, including thar associated
with low leaf nitrogen on older plants, was
directly related to stomatal conductance. Thus
stomatal conductance olfers a means for
screening for photosynthetic efficiency.
on Fischer ?t a/ (1998) suggest that remote sens-
ing that measures canopy temperature differ-
ences, which arc r€lated to the stomatal con-
ductance could be u\ed ro facilitate selecrion.
Similarly digital imaging systems could be
used lo delermrne the greenness ol ledves Ln
plots in order to select stay green" piants.
Hence not only does stay green care appear to
be an opportunity for increased yields, but
also it should be relatively simple to select for
stay grcen cane.
Hanest Index
The improvement in yield of maize and soy-
beans in the in the past fifty years has largely
been a result of increased biomass in the seed
filling period related to the stay green charac-
ter (Specht et al 1999) whereas in the small
grains yield (Evans and Fischer 1999) and
sunflower (Lopez Pereira et c/ 2000) increas-
es are largely due to increases in hafiest
index. Muchow et a/ (1996) estimated a max-
imum harvest index of 0.39 for sugarcane,
which is much lower thal that found in many
other crops. For example, the harvest index of
modem wheat varieties is about 507o and
could possibly be raised to 60% (Reynolds er
al 1999). The question then arises as to what
extent could the haflest index of sugarcane be
increased.
For the sake of argument, let us take a
one-year crop and the figure of Robertson €t
2OO1
Sugarcane growth and development
al (1999a) indicating that sralk growth and
accumulation of sucrose begins when we
have accumulated l0 t aerial biomass ha,.
The assumption is that this threshold level of
biomass is required to establish the initial
leaf area and the stalks (sinks) that will
develop during the simultaneous growth
phase. Let us then assume that we can reach
the figure of l0 t aerial biomass ha' by 160
days after planting with a leaf area index of
5. This is a very conservative estimate recog-
nizing the fact that the cane crop develops
leaf area very slowly in the early phase
growthi modem tropical rice (a C 3 plant)
varieties yield commercially 7 t paddy ha'l
with a harvest index of 5070, equivalent to 14
t ae al biomass ha" by 120 days after
planting.
If stalk growth commences 160 days after
planting, there are 200 days for growth and
accumulation of sucrose in the stalks in the
maturation phase in a one year crop cycle.
During this period we need to maintain a leaf
area index of arcund 5 (8670 interception of
the photosynthetically active radiation with
k=0.39) so as to sustain high crop growth
rates. At a specific leaf weight of 100 cm, g'
a LAI of 5 is equivalent to 5 t biomass ha'.
With a leaf longevity of 140 days (Amaya et
a1 1995) leaf area can be maintained by form-
ing l/140 of the rotal leafarea per day (*ris is
an approximation that assumes that new leaf
area is formed at a linear rate with time)
which is equivalent to about 7 t leaf hr' dur-
ing the 200 day pe od- New leaves are cost-
ly to produce in energy terms as they have
high nitrogen content and 7 t biomass ha,
would have a cost equivalent to around 10 t
of stalk biomass.
Crop growth rates of the aerial portion of
the plant can be sustained ovet prolong€d
periods at the level of about 40 g m" dayi
(Muchow er al 1994), and if the "sray green"
character is incorporatedit can be assumed
that this growth rated could be maintained
during the 200 day maturation phase. This
would give a total production of new aerial
biomass of 80 t ha-, of which
l0 t ha, biomass
equivalent are for the production of 7 t har
new leaves. This leaves 70 t ha' for stalk
growth and development.
The maximum stalk sucrose dry matter
percentage reported for whole millable stalks
is about 55% (Bull and Galzsiou 1975.
Cenicafra 2000), hence it should be possible to
obtain 70 x 0.55 = 38.5 t sucrcse ha!. This
high figure inter alia indicates that there is still
considerable scope to increase potential yield
in sugarcane.
11
Sugarca e gtowth and development
The total aerial biomass of the crop after
one year will be the l0 t ha' of initial biomass,
plus the 70 t ha'stalk biomass and 7 t ha'bio-
mass of new leaves lormed in lhe maluration
period for a total of 87 t ha'. With 38.5 t
sucrose ha' the harvest index is 4470, some-
what higher than the maximum of 39% esti-
mated by Muchow et dl 1994. lt is noteworthy
thai using the figure of maximum stalk
sucrose percentage dry matter of 487o sug-
gested by Muchow et al (1994) the estimated
sucrose yield falls to 33.6 t ha' and the haryest
index to 3970, which is precisely the same as
that estimaled by Muchou pt al t 1994)
Potential yield
Very high potential yields of 38 i sucrose ha
I
)r' appear to be wllh in thc realms of potsi
bility by combining various management and
varietal characteristics but without raising any
fundamental parameters such as RUE or stalk
sucrose dry matter content above levels readi_
ly obtained (oday LoJging resi\lanl \arrelies
with the sac) green characlet that maintain
high sucrose contents at high nitrogen levels
planted at high densities and well fertilized
should be capable of reaching these levels of
productivity.
Improviig qualil!
The quality of cane is a vital factor in deter-
mining the profitability of the sugar industry'
Two of the most important factors in deter-
mining cane quality are the sucrose content
and the ease of harvesting.
Sucfose conlent
At Cenicafia we estimate that, ceteris paribus,
a proportionately similar increase in total
sugar yield via increascd sucrose content will
give double the economic benefits of a similar
increase in cane yield. Hou ever' mo\t experi-
enced breeders and agronomists suggest that
increasing sucrose content is more difficult
than increasing cane yield lnthe Colombian
case commercial sugar yield per ha has more
than doubled in the last fifty years' whereas
the sugar recovered percent cane has
increased by less than 2070
The average sucrose contents obtained
commercially are far below those occasional-
ly obtained on an expedmental or commercial
basis on whole cane stalks, or the levels found
in the basal segmen$ of the stalks of high
yielding cane varieties (Bull and Glazsiou
1963, Lanahondo Pers. comm.). Hence it
12
would appear that in commercial production
we are way below the physiological limits for
the accumulation of sucrose on a fresh weight
basis and there are probably opportunities to
manipulate the plant at the cell and molecular
level to increase sucrose concentration
Nevertheless this article wili not dwell on the
mechanisms involved as tt falls in lhe domain
of other authors. Rather this paper will reflect
on the fact that the sugar content of present
commercial va eties are wellbelow the phys-
iologrcal limits to sugar concentration and
breeders can almoit certainly derelop van-
eties with higher sugar concentrations.
Crop husbandry and management has a
major influence on sucrose content ln most
sugarcane growing areas the harvest period is
determined by physiological response of the
crop (lnman-Bamber 1991). The harvest
operation is concentrated in the cooler and
drier periods of the year. Drying off by sus-
pending irrigation is a standard practice (Ellis
and Larkford 1990, Roberlson et al 199'/ '
1999b) and in some regions where natural
conditions do not favor high sucrose concen'
tration the crop is artificially matured with
chemical ripeners (Villegas 1995). Nitrogen
is applied early in the crop cycle and top
dressing is not pracliced. The underlying
principle behind all these practices is io stress
the crop in such a manner that growth of new
tissue is suppressed more than the rale of car_
bon assimilation.
Alrhough the pnnciple behlnd lhese van'
ous management strategies is simple, in prac-
tice it is often very difficult to achieve the
degree of stress required; too much stress will
decrease photosynlhesis to such an extent that
total yield deciines whilst insufficient stress
will result in low sugar concentrations and if
there is sufficient excess new growth may
even reduce total sugar yield. A few examples
may illustrate this pointr nitrogen supply to
thc crop cannol suddenly be turned off and
hence sufficient for optimum early growth
may be excessive in the later stages: irrigation
may be cut ofi in the drying off period but it
may rain; a warm front may increase temper-
ature and stimulate new growth in the suppos-
edly cool harvesling period.
The question then arises as to the best
means of managing these variable levels of
stress in the final stages before harvest A sim-
ilar problem occurs in the cassava crop: after
a prolonged dry period, ralnfall induces
flush of new growth and the starch content of
the roots decreases markedly and eating qual-
ity is greatly reduced. Farmers appear to have
selected varieties that do nol present such a
rapid flush of new leaves and maintain their
starch content al the onset ofthe rainy period
In sugarcane there is little solid information
on this theme, but some observations suggest
that variation may exist. Germination of the
inrtral seed sett and tnitial Srowth epilomjze
growth of new tissue: large varietal differ-
ences in germination ability have been
observed. Inlerestingly, in the Cenicaia
breeding program in Colombia, growers
rejected rhe variety with the highest and most
stable sugar content up to the moment because
ofits very slow germination. Furthermore, we
have already noted ln a different context that
some varieties appear to maintain their
sucrose content at high nilrogen levels
t suggest that it may be possible to selecl
cane varieues lhat have J lo\^er capacrly to
produce flushes of new growth towards the
end of their growth cycle and that this charac-
ter could lead to stable high sucrose content of
cane under field conditions. ln o.der to
achieve this goal breeders would have to
change their selection strategies and select for
slcrose content under conditions that are nor-
mally considered adverse for this chamcter:
that is to say selection would be made at high
nitrogen levels, with ample water and harvests
at higher temperatwes.
Ease of hah-est
The harvesting operation, including transport
to the mjll, often represents 20-307o of the
total cost of producing sugar. Futhemore,
difficulties in the harvesting operation that
result in more tmsh in cane complicate the
extraction processes in the sugar mills- Hence
there is a large incentive to imptove the ease
of harvesting and reduce costs. Obviously the
less cane that has to be harvested and trans-
ported, the lower the costs per unit sugar pro-
duced. Thus increased sucrose content is ol
paramounl impondnce in reducing harve\ling
costs, but this subject has already been
addressed in preceding sections of this docu-
ment.
In many parts of the world cane is still
manually harvested, nonetheless as countdes
dcvelop mechMical har\eshng qill increa\-
ingly become the norm. ln order to facilitate
the mechanization of the crop at the moment
ofharvest the crop should be erect to facilitate
topping and feed into the harvester, and the
a dead ard dying leaves should detach readily
from the stalks. Considerable progress has
already been made in developing vadeties that
readily shed old and senescing leaves
(Cenicana 2000). Earlier in this article the
SUGAR CANE INTERNATIONAL, AUGUST 2OO1

importance of erect cane has been shessed
and hence this characteristic appean to be of
vital importance for several reasons.
Conclusions
The yield of sugarcane has increased over
the last fifty years on a world basis, with some
countries obtaining much greater increases
I
than others. There is no evidence that average
t yields in any region aJe reachin8 a ceiling or
barrier that cannot be broken if research pro-
grams continue to develop new technology.
Although lhe basis of
many improvements
may be new varieties, their potentialwill only
be realized if they are accompanied by appro-
priate management practices.
The cane crop ol lhe lulure .hould rncor-
porate the .rldy green charactet but should
readily shed older or senescing leaves. The
crop should be erect and lodging resistant,
even when managed so as to increase total
biomass production. The varieties should
maintain high sucrose content even under
conditions that favor flushes of new
growth.
The lack of critical periods in the devel-
opmeff ofthe crop makes it relatively tolerant
of whilst the simultalreous
stress conditions.
development charactedstic of the crop
requires skilful management to achieve a del-
icate balance between overall growth and
sucrose yield.
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SUGAR CANE INTEFNAT]ONAL. AUGUST 2OO1
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. continued from page 4
Scenario Planning
This method can help avoid what Aron calls
"managerral m) opla" \\hen addressing
investments that may not show immediate
financial retums. Scenario planning lays out
a variety of paths that can occur if the
investment rs made - or tf tt i5 not made -
and pushes decision-malers to define the
likelihood for each scenario and make deci-
sions accordingly.
Real Option Theory
lmagine a game of seven-card stud poker.
On each deal, the player antes up to see his
next caJd - and his opponents'. If he doesn't
like what he sees, hc iblds. The rheory
defines technology investments as "options'
to be pursued. As opportunities present
themselves, managers can place small bets
--or investmcnts - on a variety of lT projects
and see how they play out. As one becomes
more likely to yield gains,
the betting srraF
egy" becomes more clear. "lt's not a com-
mon practice yet," Curiey says. "But it is
emerging and there will be early adopters."
Both real option rheor) and scenarro
planninB are leflile ground
for busrness
researchers and can be hrghl) esoteric.
Experts say plenty of consultants stand
ready to advise how to apply these concepts.
"R&D Council"
Curley sals Intel has pLrshed some of this
evaluation to a separate unit. devoting a
small percentage of its budget lo that unir for
research. Managers bring the more "out-
there" or innovative proposals to the council
for evaluation.
Determinitg the value depends on corpo-
rdte culture
Ultimately, business culture is critical. A
Chief Finance Officer who demands hard
numbers to justify every investment will get
what he asks for, Harker says. But he may not
be able to depend on what he gets. "People
are often looling lor a panrcea. ,ome maglc
computer program,' Harker says. "The
ansuer is usually inside thelr organrlation.
They have to have a clear procesr for decision
making and a clear articuiation of how we
account for certain costs and benefits."
(Source: Knowledge @Wharton,
http:/,ft nowledge.wharton.upenn.edu)
Sugarcane growlh and. develophent
Obituary
Anthony C. Hannah
(1946-2001)
Tony Hannah was educated in his home
country of New Zealand, where he gradu-
ated in agricultural economics. Fram
1977 Io ].97 4 he worked at Cambridge
University as a Trade Policy research fel-
low, on a project to quantiry the effects of
UK entry in the EEC.
Ftom 7974 to 1976 he worked in
Geneva as a commodity specialist with
the GATT. He joined the lS0 in 1976 as a
provisional staff member, then in 1977 as
Head of the Economics and Statistics
Division, a position he held without inter-
ruption until his death.
Tony Hannah wrote very weli and east-
ly. He was the author or co-author of many
publications, published as tSO or FAo
papers (frequently in partnership with
Kerry l\.4ulherin) or under his own signa-
ture, Quite famous were his "Worid Sugar
l\4arket Prospects", the tast one pubtished
in 1992 and covering the 90s. ln 1997,
he co-authored (with Donald Spence) a
book on "The lnternational Sugar Trade"
Moodhead & Wiley).
He was always primarily interested in
development economics and held that
consideration should be given to the
special posrtion of developing areas rn
the world, especially commodity, trade,
As such, he stuck to the commodity
agreement scheme, while .ecognizing
that it has not worked well for sugar. For
him, free trade, the best solution, stitl
had limits when applied to the develop-
ing world and he kept hitting at
''mad modellers' and "crazy free
marketers".
A very original thinker Tony Hannah
was never against a long discussion, even
straight dispute, not to say a good fight,
but was never controversial for the sake
of controversy. Above all he was totally
honest intellectually, a rare gem in the
world of economists which made him the
best of all of us sugar analysts, and
indeed, a sound, reliable and ever-
encouraging friend.
Patrick du Genestoux
July 9, 2001
15