Annals of Botany 92: 317325, 2003

doi:10.1093/aob/mcg137, available online at www.aob.oupjournals.org

Genotypic Differences in Branching Pattern and Fruiting Habit in Common

Walnut (Juglans regia L.)
A N I T A S O L A R * and F R A N C I S T A M PA R
University of Ljubljana, Biotechnical Faculty, Agronomy Department, Jamnikarjeva 101, 1111 Ljubljana, Slovenia
Received: 8 January 2003 Returned for revision: 31 March 2003 Accepted: 7 May 2003 Published electronically: 26 June 2003

Architectural analysis of 840 Slovenian walnut (Juglans regia L.) genotypes was performed to determine the
most typical and frequent morphological types and to evaluate their vegetative and generative potential. Four
branching and fruiting patterns (IIV) were detected. A 3-year-old fruiting branch, consisting of a 3-year-old
shoot plus corresponding 2-year-old and 1-year-old shoots, was used as a structural unit for quantitative analysis.
In the intermediate fruit-bearing types with mesotonic and acrotonic branching pattern (types II and III), the
total lengths of 3-, 2- and 1-year-old shoots were 385 and 380 cm, respectively, compared with 275 and 253 cm
in the terminal and lateral-fruiting types (types I and IV). In type I, 1-year-old shoots had signicantly fewer
nodes than in other types. In addition, they had a thinner basal diameter than types III and IV, and their angles
were the most erect (39 ). Only 04 out of 36 1-year-old shoots were owering with one mixed bud with 19
female owers. In type IV, 2-year-old shoots had signicantly more nodes and a larger basal diameter than
other types. One-year-old shoots in type IV are thicker than those in other types. Ratios between the number of
owering and the total number of 1-year-old shoots were 07 in type IV, 06 in type III, 05 in type II and 01 in
type I. On 1-year-old shoots in type IV, 17 mixed buds with a mean of three female inorescences per bud
were counted. Consequently, the generative potential is highest in type IV and lowest in type I. In types II and
III, growth and the ability to bear fruits are more balanced.
2003 Annals of Botany Company
Key words: Juglans regia, walnut, genotypic variation, architectural analysis, fruiting branch, generative and vegetative
potential.

INTRODUCTION
Branching and fruiting patterns are useful features for
characterizing tree canopy architecture. They are determined via tree architectural analysis, which allows quantication of the form and shape (Godin et al., 1998). Tree
shape is dened by its branching architecture (Ustin et al.,
1991), which is a major aspect of the architectural approach
to the study of plants (Guedon et al., 2001). The architectural analysis is based on three major architectural concepts:
the architectural model, the architectural unit and reiteration
(Barthelemy et al., 1991).
The architectural unit of a species represents its fundamental architectural and functional structural component. It
is composed of all categories of tree axes (Barthelemy et al.,
1991). For any tree species, there is a nite number of axis
categories, the nature and relative position of which dene
the architectural unit (Bell, 1991). The complete or partial
repetition of the architectural unit during ontogenesis is a
common phenomenon in trees (Barthelemy et al., 1991) and
is dened as reiteration (Oldeman, 1974). The result of this
process is termed a `reiterated complex'. Thus, an adult tree
is a stack of reiterations, each of which represents a
repetition of an architectural unit (Jaeger and de Reffye,
1992). The architectural model of a tree is the growth
pattern that determines its successive architectural phases
(Halle and Oldeman, 1970) and developmental sequence of
branching (Bell, 1991). According to Halle and Oldeman
(1970), who described 24 different models for tropical trees,
the architectural model is an inherent growth strategy that

denes both the manner in which the form of the plant is

elaborated and the resulting architecture. The identication
of the architectural model is based on four major groups of
simple morphological features: type of growth, branching
pattern, morphological differentiation of axes and the
position of the sex organs (Halle and Oldeman, 1970;
Halle et al., 1978).
In common walnut (Juglans regia L.), growth is rhythmic
with the axis being built up by a succession of annual shoots
(Sabatier and Barthelemy, 2001). Flowering or vegetative
annual shoots in walnut can be monocyclic, bicyclic
(Barthelemy et al., 1995; Ducousso et al., 1995; Sabatier
et al., 1995; Sabatier and Barthelemy, 2001) or even
tricyclic (Mauget, 1976; Barthelemy et al., 1995; Sabatier
and Barthelemy, 2001). Monocyclic shoots are formed by
the rst growth ush in spring. They are usually completely
preformed in winter buds. Bicyclic and tricyclic annual
shoots are also generally preformed but in intermediate
intra-annual buds (Barthelemy et al., 1995; Sabatier et al.,
1995, 1998; Sabatier and Barthelemy, 2001). On the adult
tree, monocyclic shoots are usually female, owering in the
terminal position, while the bicyclic and triyclic shoots are
mostly vegetative (Barthelemy et al., 1995).
During the juvenile period, the stem of a walnut tree has a
monopodial development (Solar, 2000; Sabatier and
Barthelemy, 2001), and is built up by the vegetative
extension of one single apical meristem (Bell, 1991).
Axillary shoots are weaker than the main axis and are in a
subordinate position (Denffer and Ziegler, 1988). As a

Annals of Botany 92/2, Annals of Botany Company 2003; all rights reserved

318

Solar and Stampar Branching and Fruiting Habit in Walnut

walnut tree ages, the development pattern changes from

monopodial to sympodial (Solar, 2000; Sabatier and
Barthelemy, 2001). This transition is linked to terminal
female owering. Sympodial axes are built up by a linear
series of shoot units. Each new distal shoot unit develops
from an axillary bud situated on the previous shoot unit
(Bell, 1991). When the axillary shoots are formed, the
terminal bud of the main axis may rest, produce a terminal
female inorescence, or die (Denffer and Ziegler, 1988).
Branching in walnut is usually proleptic (Halle et al.,
1978) with branches formed from dormant buds (Bell, 1991;
Wu and Hinckley, 2001). In some cases, the main stem
develops lateral branches in the same growing season of its
extension (Sabatier and Barthelemy, 2001). Such branches
are termed sylleptic or immediate (Caraglio and
Barthelemy, 1997), and extend simultaneously with the
apical meristem of the main stem without displaying a
resting period (Bell, 1991) and without complete bud
formation (Wu and Hinckley, 2001).
Fruiting habit is described by branching density and by
the position of owering buds on annual shoots. According
to Germain (1990, 1992), there are three types of fruitbearing habit: terminal, intermediate and lateral. Terminal
fruit-bearing types display ower buds only on terminal or
subterminal parts of the annual shoots that are growing on
3-year-old branches. Intermediate bearers display female
owers mainly on terminal and subterminal buds on the
annual shoots when they are inserted on 2-year-old
branches. Lateral fruit-bearing types display owering
buds along 1-year-old shoots. Female owering induction
affects terminal and subterminal buds and also the majority
of the axillary buds on the current growth shoot.
In the present study, genetic variability in walnut was
analysed with respect to branching and fruiting patterns.
The research represents a continuation of the previous work
started by Germain (1979) and Szentivanyi (1990), who
combined terminal fruit-bearing and late-leang French and
Hungarian walnuts with lateral-fruiting Californian cultivars, with the aim of creating late-leang and lateral fruitbearing offspring populations, which tend to be more
precocious and have a higher yield earlier than those that
bear their nuts terminally (McGranahan and Leslie, 1991).
Their work was largely based on practical experience. The
approach taken in this study is based on exact measurements
of the fruit-bearing annual shoots parameters, in order to
(a) determine the most frequent morphological types
(concerning branching and fruiting patterns) of the walnut
progenies in Slovenia, and (b) to identify the differences in
generative and vegetative potential among the branching
and fruiting types using quantitative analysis of fruiting
branches.
MATERIALS AND METHODS
The analysis included a random selection of walnut
genotypes belonging to the local population, which grows
in the south-eastern part of Slovenia, along the Sotla river
valley. The population consisted of 840 individuals
developed from randomly selected seeds, which were
taken from healthy-looking trees, originating from unknown

indigenous genotypes from nearby villages. The phenotypic

variability of the population is enormous. Due to a high
level of heterozygosity, each tree represented a unique
genotype, having grown together with others in the same
environment. The 1-year-old seedlings were planted in
19881992. The trees were not pruned, fertilized or
irrigated, and diseases, weeds and pests were not controlled.
Data collection

During the springsummer of 1998, in all genotypes a

structural unit consisting of `a 2-year-old shoot plus
corresponding 1-year-old shoots' was observed to determine its branching pattern (acrotony, mesotony, basitony).
The nature of the terminal bud on 1-year-old shoots
(owering, vegetative) was also determined. According to
Germain's (1990) classication, genotypes were divided
into terminal, intermediate and lateral types, depending on
the distribution of owering buds on the shoot, the nature of
the buds (owering, vegetative) and the age of the owering
shoot bearer. According to Caraglio et al. (1998), different
schemes of a chosen architectural type were drawn in situ.
They represented variability in branching and the fruitbearing pattern of the entire population examined.
From among the great number of architectural schemes
described, four were selected because they were the most
typical and frequent representatives of certain branching
and fruiting types in the population. They are referred to as
`morphotypes' and labelled from I to IV.
An architectural scheme was created for each morphotype (Table 1; Fig. 2). The following parameters were
dened: trunk development (monopodial, sympodial);
orientation (plagiotropy, orthotropy); growth (determinate,
indeterminate); branching pattern; and the length of the
growth unit, which is dened as the part of the annual shoot
produced during one growth ush. Out of the whole
population, 25 genotypesthe most typical representatives
of each morphotype (i.e. 100 individuals in total)were
selected.
For quantitative analysis of the walnut canopy architecture, a structural unit was dened as the `fruiting branch'
(i.e. a 3-year-old branch plus 2-year-old shoots plus 1-yearold shoots). In each tree, ve fruiting branches, symmetrically distributed in the middle of the canopy (in terms of
height) and well exposed, were selected. On hundred and
twenty-ve 3-year-old shoots were analysed with corresponding 2-year-old and 1-year-old shoots for each morphotype. The total number of shoots analysed for all
morphotypes was: 500 3-year-old shoots; 2397 2-year-old
shoots; and 1345 1-year-old shoots.
During the 1998/1999 dormant season, several measurements were performed on the selected fruiting branches on
each 3-, 2- and 1-year-old shoot unit. Three-year-old shoots
were marked as `N-2' shoots. Their length was measured.
Two-year-old shoots on the 3-year-old base were marked as
`N-1' shoots. They were measured for total number of
shoots, number of owering shoots, length, number of
nodes, branch angles and basal diameter. One-year-old
shoots on the 2-year-old base were marked as `N' shoots.
They were measured for total number of shoots, number of

M
Ov
I
No
Non
Sp
L

II

M
Ov
I
No
Non
Sp
L

III
M
Ov
I
No
Non
Sp
L

IV
S
Osv
I
No
S
Sp
L

I
S
Osv
I
No
S
Sp
L

II
S
Osv
I
No
S
Sp
L

III
S
Osv
D
F
S
Sp
L

IV

Primary branch (second order axis)

Morphotype

S
Osv
I
No
S
Sp
L

I
S
Osv
D
No
S
Sp
L

II
S
Osv
I
No
S
Sp
L

III
S
Osv
D
F
S
Sp
L

IV

Secondary branch (third order axis)

Morphotype

S
Osv
I
No
S
Sp
L

I
S
Osv
D
No
M
Sp
L

II
S
Osv
I
No
M
Sp
L

III

S
Osv
D
F
S
Sp
M

IV

3-year-old-branch (N-2)
Morphotype

Meristem potential: M, monopodial; S, sympodial.

Orientation: Ov, ortotrophy vertical; Osv, ortotrophy slanting vertical.
Growth: I, indeterminate; D, determinate.
Sexuality: NO, without owers; F, female terminal inorescence; M, male lateral inorescence.
Branching: Non, without branches; S, sparse; M, medium, D, dense.
Phylotaxis: Sp, spiral alternate.
Growth unit (a part of the shoot developed during the spring growth ush, i.e. monocyclic 1-year-old shoot): L, long; M, medium; S, short.

Meristem potential M
Orientation
Ov
Growth
I
Sexuality
No
Branching
Non
Phylotaxis
Sp
Growth unit
L

Trait

Trunk (rst order axis)

Morphotype

TA B L E 1. Architectural units of walnut morphotypes IIV

II

III
S
S S
Osv Osv Osv
I
D D
No
F F
S
D M
Sp Sp Sp
L
M ML

II

S
Osv
D
F, M
M
Sp
SM

III

S
Osv
D
F, M
S
Sp
SM

IV

Annual shoot (N)

Morphotype

S
S
S
Osv Osv Osv
D
D
D
F
F F, M
S
S
D
Sp Sp Sp
SM L M

IV

2-year-old-shoot (N-1)
Morphotype

Solar and Stampar Branching and Fruiting Habit in Walnut

319

320

Solar and Stampar Branching and Fruiting Habit in Walnut

owering shoots, length, number of nodes, branch angles,

basal diameter, number of vegetative buds, number of
mixed buds and number of female owers. Shoot length was
measured in centimetres, from the base to the top, using a
fabric tape measure. Nodes were counted from the base of a
shoot towards the topfrom the rst to the last still
distinguishable node. The angles of shoots were determined
using a special goniometer where lower values (in degrees)
represented more erect shoots. The angle of an N shoot was
represented by the value () measured between an N shoot
and an N-1 shoot, and the angle of an N-1 shoot was
represented by the value () measured between an N-1 shoot
and an N-2 shoot. The angles of three primary branches and
two secondary branches per tree were also measured. The
primary branch angle was represented by the value ()
measured between a primary branch (second order axis) and
the trunk (rst order axis), whereas the secondary branch
angle was represented by the value () measured between a
secondary branch (third order axis) and primary branch.
Total numbers of angles measured on the primary and
secondary branches were 75 and 150, respectively. The
shoot diameter was measured in millimetres at its base using
a beak-shaped measuring tool. Flowering shoots were
determined according to the presence of mixed buds (on
the terminal, subterminal or lateral position). In 1-year-old
shoots, the number of vegetative and mixed buds was
determined, as well as the number of female owers per
inorescence.
All parameters, except buds and owers, were measured
during winter dormancy. The fruit-bearing type was determined twice in a growing season: rst in winter when shoots
displayed visible scars of the previous year fruits, and then
in spring during owering.
The effects of morphotype on individual traits were
evaluated by ANOVA, and the Duncan multiple-range test
(DMRT) at P < 005 in the statistical programming package
STATISTICA for Windows (Tulsa, OK, USA, 1994).
R E SU L T S
Qualitative analysis of tree architecture: determination of
morphotypes

Passing through the orchard, 80 schemes were created in

situ after accurate observations of the trees. They represented various types of branching and fruit-bearing patterns
on the level of the structural unit `a 2-year-old shoot with
corresponding 1-year-old shoot'. Some of the schemes are
presented in Fig. 1.
Of the 80 schemes dened in the spring of 1998, four
were chosen as the most typical and frequent representatives
of certain branching and fruiting types in the whole
population. They were referred to as morphotypes and
labelled from I to IV (Fig. 2).
Morphotype I exhibits a strong acrotony. Branching
pattern is sparse and growth units are long. In addition to
monocyclic shoots, there are also bicyclic ones, and on the
branches in the canopy top there are even tricyclic
vegetative shoots. The determinate growth is typical of
annual shoots growing on 2-year-old and older shoots.

F I G . 1. The most frequent schemes of branching and fruit-bearing

patterns on the structural unit, `a 2-year-old shoot with corresponding
annual shoots' in 840 walnut genotypes of the Bistrica ob Sotli
population. Pink, 2-year-old-shoot; yellow, 1-year-old shoot; green,
current-year vegetative shoot; green with sphere, current season
owering shoot.

Flowering buds can only be found in the terminal position

on the axis of annual shoots.
Morphotype II has mesotonic branching. Branching is
more dense than in morphotype I. Growth units are of
medium length. Determinate growth can be present on
secondary branches (third order axis), and on annual
monocyclic shoots growing on 2-year-old shoots. Fruiting
shoots are in terminal and sub-terminal positions.
In morphotype III, acrotony of the shoots can be seen as
in morphotype I. However, the number of axillary shoots in
type III is higher. Branching is mesotonic on the upper half
of the shoot and growth units are medium to long. Terminal
buds on the second and third order axes often start to rest or
die. Determinate growth is developed on annual monocyclic
shoots on 2-year-old shoots. Terminal and sub-terminal
shoots bear fruits. Vegetative shoots can also be bicyclic.
Morphotype IV develops dense branching with medium
to short growth units. Primary branches (second order axis)
have a determinate growth, as well as axillary annual shoots,
on the 2- and 3-year-old shoots. Flowering buds are found in
terminal, sub-terminal and lateral positions on the annual
shoots.
Quantitative analysis of a fruiting branch

Length of monocyclic shoots. A three-year-old shoot

(N-2) is the longest (80 cm) in intermediate type III, with
acrotonic branching, and the shortest (57 cm) in lateral type
IV (Table 2). The intermediate type II with mesotonic
branching has slightly shorter N-2 shoots (79 cm) than type

Solar and Stampar Branching and Fruiting Habit in Walnut

321

F I G . 2. Architectural unit of walnut morphotypes I (terminal fruit bearing), II (intermediate fruiting with mesotonic branching), III (intermediate
fruiting with acrotonic branching) and IV (lateral fruit bearing). 1, Trunk (rst order axis) (yellow); 2, primary branch (second order axis) (turquoise);
3, secondary branch (third order axis) (light blue); 4, 3-year-old branch (N-2) (orange); 5, 2-year-old shoot (N-1) (orange); 6, 1-year-old shoot (N)
(grey); 7, current season shoot (green); 8, previous year's fruit (white circle); 9, current year's fruit (green circle). Illustration by Mitja Solar.

TA B L E 2. The mean values for monocyclic shoot length, number of nodes, shoot number, angles, basal diameter, number of
buds and inorescences, determined for different-aged shoots in four walnut morphotypes
Morphotype
Trait

Shoot type

Shoot length (cm)

N-2
N-1
N
N-1
N
N-1, total
N-1, owering
N, total
N, owering
Second order axis
Third order axis
N-1
N
N-1
N
Vegetative
Mixed
Female

Number of nodes
Shoot number

Angles ()

Basal diameter (mm)

Number of buds on 1-year-old shoot
No. of inorescences on current-year shoot

I
693a
241a
216a
72a
69a
52a
10a
36a
04a
548a
532ab
443a
387a
88a
80a
49a
10a
19a

II
788a
266a
195a
71a
71ab
76b
37b
58b
32b
582abc
515a
469ab
440bc
93a
79a
43a
11a
19a

III

IV

799a
280a
240a
70a
93c
69b
33bc
47b
30c
558ab
535ab
493b
460c
93a
84a
48a
12a
20a

571b
237a
229a
84b
91c
49ac
29c
35a
26d
614c
553b
468ab
419ab
108b
101b
31b
17b
30b

Means marked with the same letter do not differ statistically signicantly according to the Duncan multiple-range test P < 005.

III. In genotypes that belong to type I, 3-year-old bearers are

of medium length (69 cm).

Two-year-old shoots (N-1) are the longest in types III

(28 cm) and II (27 cm) (Table 2). The shortest (N-1) shoots

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Solar and Stampar Branching and Fruiting Habit in Walnut

owering N shoots is signicantly lower in type I (04).
Types III and II have 30 and 32 fruiting N shoots, and the
lateral genotypes have signicantly more fruiting 1-year-old
shoots (26).

F I G . 3. Cumulative length (cm) of 3-year-old (hatched bars), 2-year-old

(dark grey bars) and 1-year-old (light grey bars) shoots in four walnut
morphotypes.

are measured in type IV (237 cm). They are slightly longer

in type I (241 cm). Differences are not statistically
signicant.
Morphotypes do not differ signicantly in the length of
1-year-old shoots (N): the shortest N shoots are measured in
type II (19 cm) (Table 2), followed by types I (22 cm), IV
(23 cm) and III (24 cm).
The cumulative length of 3-, 2- and 1-year-old wood was
greatest in types II (387 cm) and III (380 cm), followed by
type I (275 cm). The shortest cumulative length (253 cm)
was recorded in type IV (Fig. 3).
Number of nodes. Two-year-old shoots (N-1) have more
nodes (84) in type IV (Table 2). Types III and II have 7 and
71 nodes, respectively, whereas type I has 72 nodes per
shoot.
One-year-old shoots (N) show statistically signicant
differences between morphotypes regarding the numbers of
nodes per shoot. Type I exhibits the lowest number of nodes
(69) (Table 2), this being slightly higher in type II (71). In
types IV and III, the number of nodes is signicantly higher
(91 and 93).
Total number of 2-year-old shoots (N-1) and 1-year-old
shoots (N) on the 3-year-old basis (N-2). In type IV, the
fruiting branch consists of 49 N-1 shoots, which is
signicantly less than in types II and III (respectively, 76
and 69), and similar to type I (52). Concerning the number
of N shoots, the relationship among the morphotypes is
almost the same: in types IV and I, 35 and 36 N shoots
were observed, compared with types II and III, where 58
and 47 N shoots developed on average.
Number of owering 2-year-old shoots (N-1) and 1-yearold shoots (N). The lowest number of owering (N-1) shoots
is observed in type I (10); this is signicantly lower than
that observed in other morphotypes. Type IV has 29 N-1
fruiting shoots, while in types III and II, fruits develop on
33 and 37 N-1 owering shoots (Table 2). The number of

Angles. Primary branches (second order axis) in type IV

are most widely spread (Table 2). Their angles do not differ
appreciably (Fig. 4A). In type II, the angles of primary
branches are the most heterogeneous. Angles are widest
(82 ) in type I, and narrowest (40 ) in type III. The most
erect primary branches are found in type I.
Secondary branches (third order axes) have similar angles
irrespective of the morphotype, ranging from 515 (type II)
to 553 (type IV) (Table 2; Fig. 4B). The angles in type II
are the most homogeneous. The absolute minimum is found
in type III (40 ) and the maximum (67 ) in type IV.
Two-year-old shoots (N-1) are most widely spread in type
III (493 ) (Table 2; Fig. 4C), followed by types IV and II
(468 and 469 ). The most erect shoots are found in type I
(443 ), where the absolute minimum angle is recorded
(23 ).
Angles of 1-year-old shoots (N) are wider in types III and
II (46 and 44 ) than in types IV and I (419 and 387 ;
Table 2; Fig. 4D). The homogeneity of measurements
within each morphotype is highest in type IV. Homogeneity
of measured angles is lowest in type II, in which the most
erect 1-year-old shoot was measured (82 ).
Basal diameter. Two-year-old shoots (N-1) in type IV
have a signicantly larger basal diameter (108 mm) than
other types. Types II and III have N-1 shoots with the same
diameter (93 mm) and type I has the thinnest N-1 shoots
(88 mm) (Table 2). One-year-old shoots (N) are signicantly thicker in type IV than in the other types, with the
difference ranging from 17 mm (for type III) to 21 mm (for
type II) (Table 2).
Numbers of buds and owers on 1-year-old (N) shoots

Vegetative buds. In morphotype I there are, on average,

49 vegetative buds (Table 2). Type III has 48, and type II
has 43 vegetative buds per shoot. Type IV has only 31
vegetative buds per shoot.
Mixed (owering) buds. The 1-year-old shoots of type IV
have 17 mixed buds per shoot (Table 2). Usually, there are
three owering axillary buds plus a owering terminal bud;
sometimes the terminal bud has aborted and four axillary
buds have developed fruits. Type III has 12 mixed buds per
shoot. Usually, there is the owering terminal bud plus zero
to four axillary buds. Type II has 11 mixed buds per shoot.
Terminal buds plus zero to three axillary buds bear fruits. In
type I, fruits can develop only from one (terminal) bud.
Number of female inorescences per fruiting 1-year-old
shoot

Types I and II exhibit 19 female inorescences per shoot

(Table 2). In type I, one to three inorescences are

Solar and Stampar Branching and Fruiting Habit in Walnut

323

F I G . 4. The angles of primary (A) and secondary (B) branches, and the 2-year-old (C) and 1-year-old (D) shoots in four walnut morphotypes. Large
boxes, 75% and 25%; small boxes, median; bars, non-outlier maximums and minimum values.

developed from one (terminal) bud. In type II, one to four

inorescences are developed from a terminal bud. The
highest total number of female inorescences per shoot is
eight, as the inorescences are also developed from the
owering sub-terminal bud or axillary buds. On average,
type III has two female inorescences per shoot. One or two
of these have developed from the terminal bud, but
owering sub-terminal buds and axillary buds on the
1-year-old shoot are also important. Type IV bears, on
average, three female inorescences per shoot. The highest
possible number of developed female inorescences is nine.
If the terminal bud is present, it can lead to the development
of one or two inorescences; other inorescences develop
from axillary buds. If the terminal bud has died, all female
inorescences develop from the owering axillary buds.
DIS CUS S ION
We have shown that the connection between the architecture of walnut trees and their generative and vegetative
potential can be described using architectonic decomposition and quantitative analysis of their individual constituents.
Architectural analysis of a large number of freely
growing Slovenian walnut progenies has illustrated the
great genetic variability with regard to both the topological

organization of constituents (branching pattern and the

composition of the botanical entities) and the geometrical
organization (spatial arrangement, orientation, size and
shape of constituents) of the trees.
The characteristics of the architectural unit of walnut
trees, which represent, according to Barthelemy and
Caraglio (1991), all the different categories of tree axes
and the varying ways these axes develop, have been
analysed. They are given in Table 1, together with the
architectural scheme which describes the basic structure of
Juglans regia L. A similar breakdown for Fraxinus L. is
given by Caraglio et al. (1998).
The growth unit and characteristics of 1-year-old shoots
were described. Factors that affect the number of growth
units are: genotype vigour, fruit-bearing habit, position of
the shoot inside the tree crown and the shoot angle. Three
types of shoots (monocyclic, bicyclic and tricyclic) were
found, in agreement with Mauget (1976), Barthelemy et al.
(1995), Sabatier et al. (1995), Ducousso et al. (1995) and
Sabatier and Barthelemy (2001). The majority of 1-year-old
shoots was monocyclic (formed from one growth unit).
They could be oriferous (usually on terminal position
Barthelemy et al., 1995) or vegetative. Bicyclic shoots were
found mainly in very vigorous genotypes. They were mostly
vegetative. Barthelemy et al. (1995) also report that bicyclic
shoots in walnut are usually vegetative. They were identi-

324

Solar and Stampar Branching and Fruiting Habit in Walnut

ed by the median zone of long cataphylls (i.e. scaly leaves)

connected to noticeable short internodes which separate
spring and summer growth ushes (Sabatier and
Barthelemy, 2001). Tricyclic shoots with two rings of
cataphyll scars and short internodes, which represent two
resting phases of extension, were seldom found. Two
distinctive phases, a stagnant growth phase and a vigorous
extension-growth phase, distinguished from each other by
morphological traits, were detected in the shoot growth of a
deciduous tree, Acanthopanax sciadophylloides Franch. Et
Savat. by Seino (2001). Puntieri et al. (2000) also reported
preformed and neoformed shoots in Nothofagus dombei
(Mirb.) Blume.
For quantitative research of fruit-bearing branches in
walnut trees, a 3-year-old shoot with its corresponding
2-year-old and 1-year-old shoots was chosen as a structural
unit for carrying out morphometric analyses. Shoots were
researched in detail in four different morphological types
(morphotypes). Results indicate that morphotype I exhibits
poor generative potential. On the 3-year-old branch, only
every fth 2-year-old shoot and only every tenth 1-year-old
shoot has the ability to bear fruit. There is always only one
owering bud. On average, 19 female inorescences
develop from this bud. In this morphotype, primary
branches as well as 2-year-old and 1-year-old shoots form
more erect angles than in the other three types. This
characteristic can worsen the differentiation of owering
buds in terminal fruit-bearing walnuts. On the other hand,
their growth potential is better, as demonstrated by the high
growth ratio (the ratio between shoot length and its basal
diameter) in 1-year-old shoots, and by the number of
vegetative buds on shoots, and also by the length of shoots.
The number of vegetative and fruiting shoots in walnut
was investigated previously by Ducousso et al. (1995).
Terminal, intermediate and lateral fruit-bearing cultivars of
French and American origin were included in the analyses.
It was stated, when comparing the terminal-fruiting cultivar
Soleze with the lateral-fruiting one, Lara, that the fruiting
branch in Soleze has fewer owering 1-year-old shoots
compared with Lara, while the number of vegetative shoots
is higher in Soleze. The authors concluded that terminal
fruit-bearing walnuts do not tend to have a high crop but can
regenerate very well. On the other hand, the lateral fruitbearing walnuts are more capable of owering and possess a
low ability to regenerate. The great generative potential of
the lateral-fruiting walnuts was also reported by Germain
(1990) and McGranahan and Leslie (1991).
In this study, the good fruit-bearing potential of
morphotype IV is determined by the ratio between the
number of owering shoots and the total number of shoots.
The ratios in 2-year-old and 1-year-old shoots are 059 and
074, respectively. On average, there are 17 generative buds
per 1-year-old shoot and three female inorescences per
mixed bud. In addition, lateral-fruiting genotypes have
thicker shoots than other types. Increasing thickness of
shoots is associated with increasing hydraulic conductivity
(Cochard, 1992) in various species. Thick shoots are
therefore more likely to be able to support a heavy fruit
load and to allow an appropriate nutrient and water supply
to the fruit, thus favouring higher fruit quality (Genard and

Bruchou, 1992). Lateral-fruiting walnuts are therefore

expected to have large fruits with a better kernel quality
than the fruits of other types. This conclusion differs from
that of Lauri et al. (2001), who postulated that lateralbearing walnuts show a tendency to bear nuts of decreasing
size from year to year. Nevertheless, the good generative
potential of morphotype IV is reected in the short juvenile
period. It is known that the Slovenian lateral walnuts bear
fruit in the fth or sixth year after planting, but in other
fruiting types the juvenile period can last for more than 712
years (Solar et al., 2001, 2002).
The poor vegetative potential of lateral walnuts is likely
to be attributable to the length and number of shoots, their
low growth potential, the small total number of 2-year-old
and 1-year-old shoots, and the small number of vegetative
buds on 1-year-old shoots. Three-year-old bearers are
shortest and 2-year-old shoots are shorter than those in
other morphotypes.
The growth and ability to bear fruit are more balanced in
the intermediate morphotypes II and III. In both types, 48 %
of 2-year-old shoots ower, as do 55 and 65 % of 1-year-old
shoots on fruiting branches. On average, 11 generative buds
and 19 and 20 female owers develop on 1-year-old
shoots. Both types develop 3-year-old and 2-year-old shoots
that are longer than those of morphotypes I and IV. In
2-year-old shoots, they also have longer internodes and a
better growth ratio. The acrotonic intermediate type also has
the longest 1-year-old shoots with the best ratio between
lengths and basal diameter. The great vegetative potential of
intermediate fruit-bearing walnuts can also be seen in Fig. 3.
In these morphotypes, cumulative lengths of 3-, 2- and
1-year-old shoots are 28 % bigger than in the type I and
34 % bigger when compared with the type IV. According to
Bell (1991), long shoots with long internodes and well
distributed leaves are often considered to exhibit an
exploratory capacity, i.e. extending the framework of the
plant into new territory. Such a denition can apply to
walnut with intermediate fruit-bearing type which, according to the ndings of our research, exhibit a good ability to
regenerate and self-regulate growth and regeneration.
Interestingly, the angles measured in this study indicate
that, in general, primary branches are more widely separated
than secondary ones. The 2-year-old shoots also exhibit
larger angles than the corresponding 1-year-old shoots. In
general, the present results show that higher order axes form
wider angles compared with axes of lower orders, and this is
true for all morphotypes investigated.
CONCLUSIONS
This investigation has so far enabled us to dene the
connection between the most typical morphological walnut
types and their capacity to grow and bear fruits, which
facilitates evaluation of their horticultural value. However,
to achieve a thorough understanding of this value, analysis
of the functioning of owering shoots on fruiting branches
must be carried out, extending over a period of several
successive years. In addition, research is needed to determine the genetic variability of tree structure and to study the
link between tree architecture and ecophysiological pro-

Solar and Stampar Branching and Fruiting Habit in Walnut

cesses (the inuence of light, mineral nutrition, hormones,
etc.), agrotechnical measures (pruning, bending, thinning,
etc.) and climatic conditions. Such information would be
useful in orchard management (Sabatier and Barthelemy,
2001). According to Laurens et al. (1998) and Solar et al.
(2003), long-term analysis of tree architecture and the
architecture of the fruit-bearing branch in particular, could
help to identify early morphological markers of various
phases of ontogenesis and could identify new markers for
use in walnut breeding schemes.
ACKNOWLEDGEMENT
We thank Dr Pierre Eric Lauri (Fruit Arboriculture
Laboratory, INRA Montpellier, France) for his pre-review
and comments.
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