RELATIONSIDPS BETWEEN THE VEGETATIV AND REPRODUCTIVE GROWTH OF WASIDNGTON NAVEL ORANGE CUTTINGS
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RELATIONSIDPS BETWEEN THE VEGETATIV]~
AND REPRODUCTIVE GROWTH OF
WASIDNGTON NAVEL ORANGE CUTTINGS
(CITRUS SINENSIS L. OSBECK)
ByF.LENZr
C.S.I.R.O. lrtigation Research Laboratory, Griffith, N.S.W., Australia.
SUMMARY
Experiments with rooted cuttings of Washington Navel orange, grown in
sand culture and carrying 0, I, 2 or 3 fruits, showed that increased number
of oranges resulted in reduced vegetative development, smaller fruit, thinner
skin and higher juice content. Partial defoliation reduced fruit grOwth
relatively less than vegetative growth, resulting in high fruit weight ratios.
This dominance of fruit over vegetative growth has obvious relevance to
alternate bearing in field-grown trees, but the process is studied more easily
and more completely on the cuttings. The balance between vegetative and
fruit growth affected fruit quality and such effects should always be taken
into account when interpreting environmental effects on fruit quality.
wo major problems in citrus growing are alternate bearing and fruit qUalit
Alterna:e bearing is one aspect of. fluctuation in the balance between vegetative ~
,generative growth. These fluctuatlOns are presumably mduced by climate, nutritional
status, etc. (Bouma and McIntyre, 1963). The alternate beanng 's accompanied b
"changes in fruit quality, for example smaller fruit during heavy cropping (Sham~
et at., 1933; Shamel and Pomeroy, 1934).
The effects of leaf area on the weight and volume of fruits have been demo
strated by defoliation and shading experiments with grapes (Coombe, 1959; lirova~
1958; May and Antcliff, 1963) and with pome and stone fruit (Haller and Magues '
1925; Chandler, 1957). On the other hand fruiting reduces vegetative grOwth s,
indicated by negative correlations between shoot and fruit growth in Washin:' as
o'On
. Navel orange cuttings (Gates et at., 196rb). This has been demonstrated more direct!
by Maggs (1963) who deblossomed and defruited two-year-old apple trees. On the oth;
hand, Rogers and Booth (1964) found no correlation between yield and the curren~
year's shoot growth in field-grown apple trees, but a significant negative correlatio
between yield and the following year's shoot growth. n
The present experiments with Washington Navel orange cuttings studied the
t Present address: Institut fiir Obstbau, I Berlin 33, Kiebitzweg 18-20.32 Growth of Washington Navel Orange Cuttings
eHecis of yield, defoliation and deblossoming on the development of fruit and
vegetative parts as \ve11 as on fruit quality.
lvlater£al and 1ncthods
Raising of cuttings
One \Vasllington Navel orange tree was selected and cuttings \vere taken from
the tips of spring flush shoots according to Gates et al. (rg6ra). The cuttings \\'cre
rooted in a mist propagator covered with plastic sheeting so that the temperature
was goO F. (320 C.) and the relative humidity about roo %. uniform cuttings seledcd
from this initial propagation were tmnsplantcd during June 1963 into 22' 5 em.
plastic pots filled with cleaned COa.fSC river s"ngs
ev(']opment of fruit and ~~:~~.~~~~ ~ :;r~,~~~~~'I~i~:~"~:~~I~;:;r;t~'i:~':h.~;:~~
:.;' f. L["
cuttings \vere taken from
:96ra). The cuttings Were
: so that the temperature
L'nifonn cuttings selected
fune 1963 into 22' 5 em.
nd \vas regularly leached
1'L \TE I
-~ -,4'0; K ~, 5'0; ~a-F. LENZ
33
om,
I GROWTH DRY WEIGHT AT 176 DAYS
LSD P34 Growth of Washington :Kave! Orange Cuttings
Defoliation fxp(rimcnls the orangl~5 in the defo1iat(
The effects of different degrees of defoliation on fruit development and quality of the controb.
were exalllilH~,5 F. LENZ 35
the oranges in the defoliated treatment \vere \vell developed though smaller than those
)pment and quality of the umtrol::;.
or a long period, all
Growth and gcne:ral T'SLE II
lie II. r.jfect of defoliation on mean quatity chtl}'acteristics per frllit
Fresh Vrj]umc c c c TSS Skin
OAYS ,. u
Treatment weight (ml.) '"
JUIce "
add 'ISS acid tlllckness
(gm.) (mm.)
._--
_-\11 leaves except new growth removed "5 9 25 8 3 41 - 0 o-S,~ 8'3 IO'13 0'2
Control 3 .,.,, 5 399 S 37'0 0'9 1 9'9 I r . 26 "2
140 LS.D. P < 0'05·· .. .. .. "j 64 h 78 4 ;-15 NS , '0
I
.:\.5. , .2
-------------~-----
All leaves removed, except behind fruit 144' I 1°7 -2Growth of Washington Navel Orange Cuttings
"n, GF
Defolt'atl:on excluding fruit-bearing shoots
9
\Vhen only the leaves of the fruit-bearing shoots were left, there was reduction
in fruit growth. At the end of the experiment the severity of the treatment was
demonstrated by the very small1eaf weight of the plants so treated. Considering this,
fruit growth was remarkably good (Fig. 5).
8
em. 6 '12·63
GROWTH DRY W£lGHT AT 222 DAYS
I
________ CONTROL
W 7
N
~
/ LSD. P4
~ttings F. LENZ 37
,m. GROWTH DRY WEIGHT AT 222 DAYS
left, there was reduction
,
Dn
ty of the treatment Was COI.,TP.OL
tre3.ted. Considering this,
Do
8
6 '12·63
OlGHT AT 222 DAYS
!
1
140 167·64 ~;-
6
l- FPJJIT 100 Z
~
ri.'"
5
~
~ , ,
60 50 100 ISO 200 Do Dn CONTROL
DAYS TREATMENTS
I - LEAVES
FIG. 6
Fruit size development and dry matter of plant parts in cuttings with only old cycle
leaves (Do) and young cycle leaves (Dn).
I - S1EMS 20
J - flOOTS It might be that the type of leaves could have influenced the results, but one
CONTROL
would expect less photosynthesis in the old than in the young cycle leaves. Hence the
very high fruit/plant weight ratio in the "old cycle" cuttings was all the more
TREATMENT
noteworthy.
oats, on fruit developmeniand Efficts of defoliation on fruit quality
These aIe shown in Table II. FollO\ving dcfolLation the skin thickne::;s decreased
with concurrent increases in juice %. This occurred even in the experiment illustrated
in Fig. 6, where fruit growth \vas not greatly r0'clucec1. Effects on other quality
ut 6 to 8 weeks intervals, characteristics were variable. \Vhen fruit growth was reduced there were also
:l until fruit set are called decreases in aciel antI T55, but in the experiment of Fig. 6 acid and TSS were highest
young growth cycles. The in severely defoliated plants.
of old and young growth
reatment differences were DlSCUSSIOK
with only old cycle leaves
ments. ~evertheless, fruit Fruit quality
.veights \\'ere similar in all Increase in the number of oranges per cutting increased sugar and acid contents,
1\\'th was strongly reduced and decreased skin thickness (Table I). These changes are similar to those occurring
in field-grown trees during heavy bearing.Growth of Washington Nave! Orange Cuttings
CH,"I-NDLER, \V. H.
That the balance between vegetative and gf:n~rative growth can affect fruit
CoOME.E, B. G. (1
quality was also shov,rn by the defoliation experimf'nts (Table II). Large e[[ec\:s on
aficcted by ,
frUtl qUJlity due to different l1l:.trient regimes have been reported for apples
Fitic., 10, K;
(Bunelllann, 19(4) and for citrus (Parker and Jones, 195T), but whether these were
GATl'S, C. T., Bor!»
all direct effects O~ were partly due to changes 1:1 the balance hehvee-:-t vegetative allll
of the \Vasb
generative growth is not kr:own. Thus interpretation of s'tlch stud~es would be
of the rootf'.(
facilitated if plants Wit11 equal numbers of fruits were used.
GATES, C. T., BOl
cntti'!.1gs of
Fruit and vegetat1've growth
frliting cu:1
A strong dominance of frnit over vegetative development was shown ill th2 H ALl.ER, :'it H., ar
preseut experiments with \Vashington Navel orange::;. Tncreased number of (rnit~ and compos
reduced vegetative devdopment. :Moreovcr reduction in le;:d area, due to GckliaLiou HOAGLAND, D. R.,
4 weeks after fruit set, had a stronger depressing effect on Vf~getative than on fruit plants with(
developmcnt, the latter not being reduced in one instance c1('spite a reduction iE KInD, F., and WE'
foliage to one-fJ!th of the control (Fig. 6). Thus suggestions that the fmit is dominant fruits after
in t11C developmcnt of citrus (Gates et al., 196Ib) as it is in other :::pecies (\Vi1cox, 1937; LEONAR", E. R. (
Leonard, 1962) were confirmed. The strong fruit developner.t, even in plants with a special refel
very small leaf area (Fig. 5) suggests high net assimilation rates, which would be MAGGS, D. H. (I<
similar to increases in photosynthetic efficiency in fruit-bearing trees of peach ~nd fruiting. J.
apple (Ryugo and Davis, "959; Mochizuki, J962; J\lagg3, 1063). MAY, P., and AN~
Apart from the eompetiLiu!l between vegetative organs and the fruit then'. \.\/;1.S. in the Sult'
also competition between indiviuual fruits on the same plant lFig. I). MOCHIZUKI, T. (I'
Dominance of the fruit is presumably res?onsible for the induction of altrrnate apI)le tree \
bearing in fielrl-grmvn trees. F::uiting leads to depletion of reserves, as is demon ~, 4 0 - 124 .
strated by the red11ced de\.cclopmcnt of root and ste,n in heavily bearing plants NOVAK, ].(195 8',.
(Fig. I), and partic.nlarly after defoliation (Fig, 3). of gral)cs, ,
It is not kno'Nn ,,,,hich metabolites or Illinc-:'"al nutrients become limiting, nor how PAHRER, E. R., a
tlw successful compf'.tition of the fruit over the vegetative organs comes about. and lJu(lJil~
Future \vark on this shnl1kl be initiated on cutCngs, where all plant parts can be RUGEKS, VI/T.S., a
quantitatively studied, rather than on ficld-growll trees. apple.J. h,
RVL:Go, I{., and I
I wish to thank 11r. E. Hoare, Officer i'l Cha::ge C.S.I.1\.. O. IrngaTion Research , rate of pea
Laboratory, Griffith, N.S.\V., for his kind support of Lllis work and 1 gladly acknO\v PrOG. A 11;0
ledge my indebtedness to lIdr, G, 'kIntyre for :::..tatistical treatment of the data, to SI1A~1EL, A. D., ~
.\liss ~\i. Stokes, 11r. H. Gilliard and ~'IL T. l'IIitchel for able ter.Jwical assistance, ar.d to size in Val
Dr. H. Greenway and Mr. R. de Fossard for critical suggestions wltile pre?aringthe paper. SHAME:", A. D., I
\ size in \\1::;
EEFEHENCES "hecox, J. C. (,
between f
BOl'MA, D., and. :l\lcINTYRE, G. .i\.. (1963). A factorial field exp~riment with citrl1S. I
j. hort. Sci., 38, I75-98.
B(~.\"EM . \NN, G. (I90.1). Die Fruchtqualitat bc:m Apfel in Abhangigkeit yon der
i\Tabrstofizu:uhr. f. Litera:ur-Dbersicbt (Samrr:elrde:-atl. GartenbaU1o£ss., 29, \
:f- Sr-5!6.""o,L',~"',=0_.- - - - - - - - - - - - 0"
tings F. LENZ 39
rO\vth can affect fruit CH.-\~DLER, \V. H. (1957). Deciduous orchards. H, Eimpton, London, 3rd edition.
c II), Large effects on COO;,[BE, B. G. (1959). Fruit set and development in seeded grape yariettes as
1 reported for apples affected by defoliation, topping, girdling and other treatments. Am. j. Enol.
ut whether these were Vitie., 10, 85-IOO.
lehveen vegetative and G_\TES, C. T., BOL"}lA, D., and GROE:K'E\VEGEK, H. (I96Ia). The development of cuttings
uch studies would be of the "\Vashington :Kavel orange to the stage of fruit set. 1. The development
of the rooted cutting. Aust. j. agric. Res., 12, 1050-65.
GATES. C. T., BOU}lA, D., and GROEKEWEGE~, H. (rg6rb). The development of
cuttings of the \Vashington Navel orange to the stage of fruit set. III. The
fruiting cutting. Aust. j. agric. Res., 12, IOS1-8.
ent was shovm in the
HALLER, M. R, and MAGNESS, J R. (1925). The relation of leaf area to the growth
ased number of fruits
and composition of apples. Froc. Amer. Soc. hort. Sci., 22, 189---g6.
lrea, due to defoliation
HOAGLAND, D. R. and ARKON, D. I. (1950). The water-culture method for growing
2getative than on fruit
plants without soiL Cire. CatiI agrie. Exp. Stn., No. 347.
despite a reduction in
KIDD, F., and WEST, C. (1947i. A note on the assimilation of carbon dioxide by apple
it the fruit is dominant
fruits after gathering. Nc'''' Phytol., 46, 274-5.
r species (Wilcox, 1937;
.J even in plants with a
LEONARD, E. R. (1962). Inter-relations of vegetative and reproductive growth, with
rates, which would be special reference to indeterminate plants. Bot. Rev., 28, 353-410.
MAGGS, D. R (1963). The reduction in growth of apple trees brought about by
ing trc('~ of peach and
fruiting. I IIort. Sci., 38, II9-28.
).
md the fruit there was MAY, P., and ANTCLlFF, A. J (1963). The effect of shading on fruitfulness and yield
(Fig, 1), in the Sultana. J- hort. Sci., 38, 85-94
ivloCHIZGKI, T. (I962). Studies on the elucidation of factors affecting the decline in
; induction of alternate
apple tree vigour as induced by fruit load. Bull. Fac. Agric. Hirosaki Univ. No.
reserves, as is demon
8, 40-I24.
heavily bearing plants
NOV,\K, J. (1958). The effect of reduced leaf surface of vines on quantity and quality
of grapes. Arh. poljopr. Nauk., 11. 82-92; Hort. Abstr., 29, 2242, 1959.
~come limiting, nor how
PARKER, E. R, and Jo,ms, W. W. (1951). Effects of fertilizers upon the yields, size
e organs comes about.
and quality of orange fruits. Bull. Calif. agr. Exp. Stn., No. 722.
all plant parts can be
ROGERS, \V. S., and BOOTH, G. A. (1964). Relationship of crop and shoot growth in
apple. I hort. Sci., 39, 61-5.
RVUGo, K., and DAVIS, L. D. (1959). The comparison behveen the net assimilation
.0. Irrigation Research
rate of peach leaves and the rate of accumulation of dry weight by the crop.
'k and I gladly acknow
,atment of the data, to Proc. Amer. Soc. hort. Sci., 74, 134-43.
SHAMEL, A. D., and POMEROY, C. S. (1934). Relation of amount of foliage to fruit
mical assistance, and to
'hile preparing the paper. size in Valencia oranges. Calif Citrog., 19, 140-1.
SHAMEL, A. D., POMEROY, C. S., and CARYL, R E. (1933). Relation of foliage to fruit
size in \Vashington Navel oranges. Calif. Citrog., 1S, 296.
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Abhangigkeit von cler
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