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Amino acids in kiwifruit 1. Distribution

within the fruitduring fruit maturation
E.A. MacRae
a

a b

& R.J. Redgwell

a b

DSIR Fruit and Trees , Private Bag, Auckland , New Zealand

The Horticulture and Food Research, Institute of New Zealand

Ltd , Private Bag 92169, Auckland , New Zealand
Published online: 05 Jan 2012.

To cite this article: E.A. MacRae & R.J. Redgwell (1992) Amino acids in kiwifruit 1. Distribution
within the fruitduring fruit maturation, New Zealand Journal of Crop and Horticultural Science, 20:3,
329-336, DOI: 10.1080/01140671.1992.10421775
To link to this article: http://dx.doi.org/10.1080/01140671.1992.10421775

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New Zealand Journal ojCrop andHorticultural Science, 1992, Vol. 20:329-336

0114-067119212003-0329 $2.50/0

329

The Royal Society of New Zealand 1992

Amino acids in kiwifruit

1. Distribution within the fruit during fruit maturation
E. A. MacRAE*
R.J.~VVE[JL*

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DSIR Fruit and Trees

Private Bag
Auckland, New Zealand
*Present address: The Horticulture and Food Research
Institute of New Zealand Ltd, Private Bag 92169,
Auckland, New Zealand.
Abstract The distribution of free amino acids in
different tissues ofkiwifrnit (Actinidia deliciosa) and
changes in their concentration during maturation of
the fruit were investigated. Amino acid concentrations
in the fruit (and each fruit tissue except the skin)
decreased as the fruit matured. Asparagine and
glutamine were the main amino acids to decrease,
allowing arginine and y-aminobutyric acid to become
the predominant amino acids in fruit harvested at the
end ofMay. There were differences in individual and
total amino acid concentrations between the core,
inner and outer cortex fruit tissues. and with the fruit
stalk. The core contained the highest concentration of
amino acids and the outer cortex the least In February,
asparagine was the major amino acid in the inner and
outer cortex, whereas arginine and y-aminobutyric
acid were predominant in the core. By May, arginine
and y-aminobutyric acid were predominant in all
tissues. Arginine and/or y-aminobutyric acid always
formed a longitudinal gradient within the fruit,
regardless of tissue type. It was highest at the stem
end and lowest at the blossom (distal) end.
Concentrations of amino acids in the fruit stalk
increased as the fruit matured. Arginine and/or yaminobutyric acid and asparagine were the major
amino acids to increase, followed by glutamine.

H91093
Received11 November 1991; accepted 26 May 1992

Keywords Actinidia deliciosa; kiwifruit; cv.

Hayward; arginine; asparagine; glutamine; maturation

INTRODUCTION
Previous investigations of the chemical constituents
ofkiwifrnit (Aetinidia deliciosa (A. Chev.) CF. Liang
et A.R. Ferguson) have indicated that the different
tissues vary in chemical composition, e.g., organic
acid and starch (MacRae et al. 1989a). Calcium
concentrations have also been found to vary within
the fruit longitudinally (Ferguson 1980), and were
highest in the inner cortex and at the stem end of the
fruit. As well, some events in fruit ripening such as
cell wall and starch degradation and softening do not
occur synchronously in all tissues (MacRae et al.
1989b; Redgwell et al. 1990; Hallett et al. 1992). In
these investigations no measurement was made of
free amino acids in fruit. Fuke & Matsuoka (1982)
found little change in total free amino acids during
kiwifruit growth and in fruit held for 1 month after
harvest. In their experiments arginine was the
predominant amino acid in fruit and decreased as
fruit matured and ripened. Alanine content ina-eased
with ripening. Clark et aI. (1992) found that the
concentration of glutamine and asparagine ina-eased
markedly early in fruit development, peaking at 9
weeks after fruit set. After this time both amino acids
declined 2-3-fold as fruit continued to develop and
mature, and the level ofseveral amino acids increased
during low temperature storage. Arginine is regarded
as a key amino acid in terms of nitrogen (N) storage,
and Clark et al. (1992) found a strong correlation
between arginine concentrations in the fruit and N
fertiliser regimes. Increased N resulted in increased
arginine concentrations.
In experiments on the fate of l4C-leaf photosynthate in fruit during fruit maturation (MacRae &
Redgwell 1990), decreased proportions of 14C_
photosynthate were allocated to the amino acids as
fruit matured. Except for fruit in February, the
proportion of label found in amino acids also decreased
with time after application of radioactivity. However,

New ZealandJournalof Crop and Horticultural Science,1992, Vol. 20

330

increased allocation of 14C-photosynthate to amino

acids in the fruit stalk occurred as the fruit matured.
This may well be reflected in amino acid concentrations,and indicatea major shift in requirementfor
aminoacids,a factornot found in concurrentanalysis
of leaf petioles.
We describe in this study the composition and
distribution of amino acids in the fruit and fruit stalk
at three stages of fruit maturation.
MATERIALS AND METHODS

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Plant material

Kiwifruitwere harvestedin 1986from a l();oyear-old

vinegrowingina blockat theDSIRResearch Orchard
at Kumeu, New Zealand as previously outlined
(MacRae & Redgwell 1990). On each sampling
occassion, fruit were harvested from two lateralson
opposingsides of the vine; four fruit per lateral;with
4-6 terminal leaves on each lateral.The experiment
tookplace at threestagesof fruitdevelopment: in late
February wben fruit were 55% of ftnal fresh weight,
bad unripe seeds, and a soluble solids concentration
00.9%; late April whenfruitwere82%offmal fresh
weight, seeds were mature, and fruit bad a soluble
solids concentration of 5.4%; and late May wben
fruit had a solublesolidsconcentration of9.2%. Fruit
set occurredin early Decemberand furtherdetailsof
fruitdevelopment arereported inMacRae& Redgwell
(1990). Nitrogen as urea. totalling 140 kglha, was
appliedin earlyspring(August)and beforeflowering
(late October)in a ratio of 2:1.
The water solublefractionfrom differentpartsof
eachfruit(fruitstalkand skin; blossom(distal), middle

and stem ends of the outer cortex, inner cortex, and

core)was extractedas previouslydescribed(MacRae
& Redgwell 1990), and the amino acid fraction
isolated.
Quantitation of free amino acids

Amino acids were analysed as their phenyl

isothiocyanate derivatives by highperformance liquid
chromatography (HPLC) (Heinrikson & Meredith
1984). A set of standards were co-derivatised with
representative kiwifruit amino acids to check
derivatisation efftciencies. Recoveryof eacb always
exceeded 95%. y-aminobutyric acid (GABA) was
not quantifted by this technique as it co-chromatographed witharginine; and arginineconcentrations
reported will be an overestimate and are written
arginine/GABA. Estimates from thin layer
chromatograms (Bieleski & Turner 1966) indicated
that arginine was present in greater quantities in
componentfruit tissues than GABA. This was in the
order of 100:1 in the core, 10:1 in the inner cortex,
and 2:1 for theoutercortex.Whenadjustedfor whole
fruit (using 7.9% for core, 35.8% for inner cortex,
and 53.6% for outer cortex as relative tissue
contributions) a value of 3:1 is obtained. This is
similar to the value reported by Clark et al. (1992):
c. 2.5:1 for whole fruit.
Methionine was measured but has not been
reported because of variability in quantitation,
probably because of its known instability and rapid
oxidation during most extraction procedures.
Methionine concentrations ranged from 0.01 to
0.2 ~g1g fwt which representedfrom 0.1 to 1.6% of
the total amino acids.

Table 1 Concentration of the total amino acids in kiwifruit fruit and fruit stalk during the later stages
of fruit maturation. Whole fruit values were calculated using the formula of core = 7.9 %fwt, inner
cortex =35.8 %fwt, and outer cortex =53.6% fwt (see MacRae et aI. 1989a). The skin was not included.
A number of harvest parameters are included in the Table heading for reference. These are date or mean
fruit measurements from eight replicates per harvest.
Total amino acid (Ilg!g fwt)

Tissue

Harvest date:
Stalk (g fwt):
Fruit (g fwt):
Soluble solids(%):

Feb
0.59
63
3.9

Apr
0.67
94

5.4

May
0.73

115

9.2

Fruit stalk
457
4491 P < 0.001
6915P<0.001
Fruit skin
195
174
159
Whole fruit
484
154
208
3152 P < 0.001
Core
767
1127 P<0.05
Inner cortex
455 P<O.ool
217P< 0.001
173
Outer cortex
128 P<O,ool
72P<O.ool
55
_ _ _ _ _ _ _ _ _ _.::c::..:_
_-'----'--- _ _-:..=--'-_--'-=---_
_-=...:..
_

331

MacRae & Redgwell-Amino acids in kiwifruit 1. Distributionduring fruit maturation

4000

3500 -f

--...
c:
0

co

c:

oQ) _~

3000

II

Feb

Apr

May

2500

c:
0
C)
o ....... 2000

"'0

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-0co

C)

::L

1500

c:

1000
500
0
ASP

GLU

ASN

SER

GLN

HIS ARG/GABA THR

Amino acid
Fig.l Concentration of amino acid in the fruit stalk in February, April, and May. Values are means SEM of eight
replicates, and annotated with the standard amino acid abbreviations.

Statistics

Data fromtheeight singlefruitreplicates weremeaned

and either presented withSEM, ocwithlevelsof significance calculated using the standard Students r-test,
RESULTS AND DISCUSSION
Amino acids of the whole fruit and changes
during fruit maturation

The total concentration of amino acids in each tissue

of the fruit decreased as the fruit matured (fable 1).
Although there were changes in concentrations of
severalamino acids these weremostly tissuespecific.
The relative contribution of each tissue remained
similar at each stage of maturation; with the core
showing the most variation, from 5% in February to
7.9% in May.
In the whole fruit, asparagine and glutamine
concentrations decreased as fruit matured (fable 2).

Table 2
Changes in concentration of asparagine,
glutamine, and arginine in the fruit during fruit maturation.
Whole fruit values were calculated as for Table 1.
Apr

May

Amino acid concentration (Jlglgfwt)

asparagine
116.7 P < 0.001
glutamine
113.8 P < 0.001
arginineiGABA
140.7 P < 0.05

6.7
29.7
80.6

12.7
76.9

Amino acid proportion (%)

asparagine
25
glutamine
18
arginineiGABA
20
histidine
0.8

4
16
29
1.4

1
9
36
2.1

Feb

1.7

The decrease in asparagine was found in each tissue.

Histidine, isoleucine, and phenylalanine remained
similar (data not shown). The proportional
contribution of histidine and arginine/GABA to the

New ZealandJournalof Crop and Horticultural Science, 1992,Vol. 20

332

50

ASP

GLU

Stem

Blossom

fZI

c:

-co

....

40

30

ASN

GLN

ARG/GABA

Middle

c: _

oQ) -~
oc: -0)
0

......

"'C

0)

- _
:::L
0

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co

20 -

c:

10

o
Feb Apr May

Feb Apr May

...

Feb Apr May

h
Feb Apr May

n
Feb Apr May

Month
Fig.2 Amino acid concentrations in the outer cortex of the fruit in February, April, and May. Values are means SEM
of eight replicates, and annotated with the standard amino acid abbreviations.

total amino acids increased, and that of asparagine

and glutaminedecreased (fable 2).
The asparagine changes appear to be a genuine
loss from the fruit and not a dilutioneffect. If values
are estimated on a per fruit basis, as outlined above
and using the fruit weights in Table I, there was
7350 Jlg asparagine per fruit in February. In April
this reduced to 630 ug and in May to 195 Jlg.
These results conflict with those of Fuke &
Matsuoka(1982). Theyfoundlowlevelsof asparagine
(-8 Jlg/g fwt) throughoutfruit maturationand ripening, implyingan increasein totalasparagineper fruit
as fruit matured. Their value is similar to our April
sample and perhaps reflects a poor supply of N,
differentfertiliserregimes, or a non-reducing source
of N. Their total amino acid concentrations were,
however, slightly higher than ours (~ Jlg/g
fwt). The values of Clark et al. (1992) for individual
amino acids such as asparagine and glutamine are

similar to or higher than ours. The trends with fruit

development are similar, but are more dramatic in
our study. For example, in Clark et al. (1992)
asparagine decreased from c. 3 to 1.25 umol/g fwt,
whereas in the current study asparagine decreased
from c. 0.8 to 0.01 umol/g fwt. A key difference
between the two studies is the N fertiliser regime,
and the way in which the amino acids were
extracted and analysed e.g., fresh tissue versus dry
tissue.
Valuesfor N in kiwifruitin Apriland May can be
found in the literature. Ferguson & Eiseman (1983)
reporteda valueof 0.53 mg N/g fwt for fruitin April!
May. Clark & Smith found values of 0.51-0.59 mg
N/g fwt at a similar time of the season (1988) and
1.42-3.3 mg N/g fwt depending on the stage of fruit
growth(1991),and Prasadet al. (1984)0.24-0.71 mg
N/g fwt. In our experiment, total amino acid in
February was 0.48 mg/g fwt and in April 0.21 mglg

333

MacRae & Redgwell-Amino acids in kiwifruit 1. Distribution during fruit maturation

ASP

200

c:

,....0IV
....'-c:
Q)

c:
0

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"0
0
IV

150 -

GLU

Stem

IZI

Middle

ASN

GLN

ARG/GABA

Blossom

....
~

C)

.......

C)

::L.

100

c:

50 -

In_

o
Feb Apr May

Feb Apr May

Feb Apr May

I m
Feb Apr May

m
Feb Apr May

Month
Fig.3 Amino acid concentrations in the inner cortex of the fruit in February, April, and May. Values are means SEM
of eight replicates, and annotated with the standard amino acid abbreviations.

fwt and May 0.15 mglg fwt When amino acid is

calculated in terms of N, in February a value of
102 ug N/g fwt and in Maya value of 36 ~g1g fwt is
obtained for fruit N in the form of free amino acids.
When these are compared to reported values 20% of
fruit N occurred as free amino acids in February and
7% in May. These values are lower than those reported
by Clark et al. (1992), and may reflect differences in
N fertiliser regimes, all of which appear to be higher
than that in our study. An earlier study by Clark &
Smith (1991) found that free amino acids contributed to between 7 and 43% of total fruit N, more
similar to our experiment In their study vines had
one application ofN as urea (480 kg/ha) in September,
but at a higher application rate than in our
investigation.
,
In our study atginine/GABA could have
accounted for c. 5% of fruit N in May. In that of
Clark & Smith (1991) arginine/GAB A was 50% of

fruit amino acid concentration at commercial maturity

in May and hence probably contributed c. 3.5-4% of
total fruit N.

Amino acids of the fruit stalk and changes

during fruit maturation
Total amino acid concentrations of the fruit stalk
increased as fruit matured (Table 1). The major amino
acids in the stalk were arginine!GABA, glutamic
acid, glutamine, and asparagine (Fig. 1). Arginine!
GABA increased from 19% of the total amino acids
in February to 50% in May, and asparagine from 7%
in February to 29% in May. Glutamic acid decreased
from 25% in February to 4% in May and glutamine
from 15% in February to 8% in May.
All other amino acids increased in concentration
on a fresh weight basis during fruit maturation;
although their proportional contribution to the total
amino acid pool decreased. The exceptions were

New ZealandJournalof Crop and Horticultural Science, 1992,Vol. 20

334

2000

--

ASP

ASN

GLU

ARG/GABA

GLN

1750

c::::

1500

l
~

c:::: _

Q)
o

-3=

c:::: o
en
o .......

"'0

<3

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en

:::1.

1250

Blossom

IZI

Middle

Stem

1000
750

c::::

500
250

,.~ I - , ' I . , ' I

Feb Apr May

.i" .,'

I -,"

Feb Apr May

i'"

'Ipe

0,

Feb Apr May

'111'

_I' I

lip

Feb Apr May

i . " ! - .. I .~ J

Feb Apr May

Month
Fig. 4 Amino acid concentrations in the core of the fruit in February, April, and May. Values are means SEM of eight
replicates, and annotated with the standard amino acid abbreviations.

threonine (2.5%), phenylalanine (0.7%), and

isoleucine(0.8%).
The most striking feature of these results is the
change in the contribution of the fruit and fruit stalk
as comparative sources of free amino acids. In
February 99% of the free amino acids were found in
the fruit; in May only 75% werefoundin the fruit (on
a totalfresh weightbasis).Thisrepresentsan increase
of 24% in the stalk and is an enormous contribution
to the free amino acid pool by a tissue contributing
< 1% of the total fresh weight of fruit and stalk. The
amino acids that contribute to the increase in amino
acid concentration of the stalk are those that are
important in terms of N storage or transportarginine,asparagine, and glutamine(Fig. 1).
Amino acids of the different fruit tissues and
changes during fruit maturation

The outer cortex had the lowest concentration of

amino acids and the core the highest(fable 1). In all

three tissues asparagine and glutamine decreased as

the fruitmatured (Figs2, 3,4). In the inner and outer
cortexarginine/GABA decreased at theblossomend
and in the middle of the fruit during maturation, but
increased at the stem end. ArgininelGABA always
decreased in the core. In all three tissues, however,
there was a strong longitudinal differential in the
concentration of arginine/GAB A. ArgininelGABA
was always highest at the stem end of the fruit,
especiallyin April and May.
Asparagine, glutamine, and arginine/GAB A were
the predominantamino acids in all tissues; 89% in
the core, 73% in the inner cortex, and 53% in the
outer cortex in February. By May these values had
decreased to 66% in the core, 50% in the inner
cortex, and 41% in the outer cortex. The two other
major amino acids of the outer cortex were aspartic
acid and glutamic acid. Together they contributed
40% of the total in Februaryand 48% in May. In the
core they were 7% of the total in February and 27%

Downloaded by [5.12.24.26] at 13:47 20 April 2015

MacRae & Redgwell-Amino acids in kiwifruit 1.Distributionduring fruit maturation

in May; in the inner cortextheywere 20% in February
and 38% in May.
Concentrations of isoleucine, leucine, and
phenylalanine remained similar in all tissues at all
stages of fruit maturation (data not shown). Valine,
alanine, and threonine decreased slightly as fruit
matured. Concentrations of all these amino acids
were least in the outer cortex and greatest in the core.
The inner cortex differedfrom the other tissuesin
that high concentrations of aspartic and glutamic
acids were found at the blossom (distal) end of the
fruit Glutamine tended to be similar. This tissue
contained seeds and the vasculartissue feeding them.
In the skin of the fruit asparagine was the only
amino acid to show significant loss during fruit
maturation (data not shown), and together with
glutamic acid (30%), arginine!GABA (25%), and
aspartic acid (20%) formed c. 75% of the total amino
acid pool.
Tadeo et al. (1988) found that asparagine
decreased in the rind of navel orange duringripening,
but not the juice sacs. In the rind it was the major
amino acid. Arginine increasedin both rind andjuice
sacs, although it was not the predominant form of
amino acid in ripe fruit. Their findings implied that
asparagine in the fruit was continuallybeing utilised,
because asparagine was lost at a time when there was
expected to be reduced synthesis in the roots and
transport to the fruit. The same may well be true for
kiwifruit, in that both fruits are autumn/winter
harvestedand both exhibit a major lossof asparagine.
The increase found in asparagine in the fruit stalk
in kiwifruit in April and May also implies however,
that there may be some impediment to transfer of
asparagine to the fruit, or that the stalk has a stronger
requirementfor asparagine than the fruit.
The unusualgradientfoundin thefruitforarginine!
GABA is difficultto interpretwithcurrentknowledge.
It is the reverse of that found for sugar concentrations
where % soluble solids is highest at the blossom
(distal) end of the fruit in April and May (Hopkirk et
aI. 1986; MacRae et aI. 1989a). A possibility is that
there is limited movement of arginine once in the
fruit. Because there are such large quantities of
arginine in the fruit stalk in April and May it is
possible that the high concentrationsin the stem end
of the fruit are a "spill over" from the stalk.
Both the explanations for the patterns found for
asparagine and arginine presume that they are
translocated intact into the fruit and not synthesised
in situ.Examinationofthexylemsap showsglutamine
to be predominant early in fruit growth (Clark &
Smith 1991),but arginine,asparticacid,and glutamic

335

acid may be found in similar quantities in April or

May. As arginineisnotbelievedtobe a goodcandidate
for translocation, examination of phloem exudate
andlabelling studiesshouldhelp toclarify thequestion
of synthesis in situ versus translocation.
ACKNOWLEDGMENTS
We thank Noel Turner, Ross Ferguson, and Chris Clark
for helpful discussions. The New Zealand Kiwifruit
Marketing Board provided partial funding for this work.

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estimation of amino acids in crude plant extracts
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phytologist 108: 399-409.
----1991: Seasonal variation of nitrogenous
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Annals ofbotany 68: 441-450.
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