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From Tree Architecture To Tree Training - An Overview of Recent Concepts Developed in Apple in France

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From Tree Architecture to Tree Training – An Overview of Recent Concepts


Developed in Apple in France

Article · January 2002

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J. Kor. Soc. Hort. Sci. 43(6):782−788. 2002.

From Tree Architecture to Tree Training – An Overview of


Recent Concepts Developed in Apple in France
Pierre-Éric Lauri
Équipe Architecture et Fonctionnement des Espèces Fruitières, UMR BDPPC, Institut National de la
Recherche Agronomique (INRA), 34060 Montpellier cedex 01, France

Abstract. Since four decades research on training systems at INRA (Institut National de la Recherche Agronomique)-France
has focused on knowledge of fruit tree growth and fruiting patterns as a mean to improve training concepts and tools. The
underlying principles of the training concepts developed nowadays arise from two initially independent and now merged
streams of research. The first one is plant architecture (sensu Hallé et al., 1978) which developed general concepts on
growth, branching and fruiting within the entire plant kingdom, e.g. architectural model and reiteration. The other one is
the apple tree typology (sensu Lespinasse and Delort, 1986) with specific insights into vegetative growth-fruiting relation-
ships. This article reviews (i) the main research concepts developed in both areas and (ii) training considerations which stem
from them. Two examples, taken from the recent history of training systems successively developed in France, will illustrate
some of the architectural rules used to improve procedures better suited to the fruit tree biology. The first one is the transition
from renewal pruning (Vertical Axis) to the free growing fruiting branch (Solaxe), to better control the balance between veg-
etative growth and fruiting. The second one is the use of the extinction procedure (Centrifugal training concept) to optimize
light distribution in the tree, fruit color and return-bloom.
Additional key words: fruiting strategies, extinction, Solaxe, Centrifugal training system

Introduction this paper opens the way to training methods based on a more
accurate knowledge of morphological and functional character-
Two contrasting approaches to the fruit tree are generally istics of cultivars.
found in the literature on tree and orchard management. In the The underlying principles of the training concepts developed
first approach, referred to in this paper as the “physical nowadays at INRA -France stem from two initially independent
approach”, often focusing on light climate in relation to total and now merged streams of research, one on plant architecture
yield and productivity, the tree is seen as a black box. In this with an initial focus on forest trees, the other on apple tree
case, only external features such as trunk and crown diameter, typology with specific insights into vegetative growth-fruiting
tree height or leaf area index (LAI) are required and structural relationships.
components and internal organization of the canopy, e.g. distri- In this paper I will review (i) the main research concepts
bution in space of the various shoot types, are poorly known. developed in both areas and (ii) training considerations which
In this approach, canopy structure is described by statistical dis- stem from them. Since practical considerations have been
tributions (Sinoquet et al., 1993). Simple geometrical models detailed in various articles, only underlying principles are pre-
are generally assumed, e.g. cylindrical or conical shapes of can- sented here.
opy for individual trees, or a rectangular shape for continuous The research field covered by this subject is broad, with var-
hedgerows (Gussakovsky and Shahak, 1999). In the second ious topics addressed. Relevant references to the literature, in
case, the “biological approach”, the tree is seen as a highly inte- English whenever possible, have been selected if more detailed
grated living organism whose detailed growth, branching and approaches are required.
fruiting characteristics are known (Costes et al., 1997).
These two viewpoints, stemming from two quite opposite Fundamentals of Tree Constructional
visions of the tree have practical consequences for tree manip- Organization
ulation. The physical approach typically leads to the manage-
ment of tree shape through mechanical practices as hedging and Tree architecture: architectural model and reiteration. The
topping. The effects of such practices depend on many factors exuberant world vegetation, especially in the Tropics, has for a
such as plant material (the increased shading usually observed long time prevented researchers from gaining a comprehensive
has negative effects on red-colored cultivars) and amount of knowledge of whole-plant development. Classical botanical
regrowth. Past experience generally led to the conclusion that studies were developed with deep insight into the morphology
mechanical pruning results in excessive well-illuminated vege- and anatomy of specific morphological categories such as
tative growth and, at the same time, poorly developed fruiting leaves, inflorescence or flowers and fruit (Bell, 1991) but with
shoots (Jackson, 1980). a lack of integrated knowledge on whole-plant development.
In contrast, the biological approach that will be presented in The quest for knowledge of general underlying rules of plant

Received for publication 11 October 2002. Accepted for publication 18 November 2002.

782
J. Kor. Soc. Hort. Sci. 43(6):782−788. 2002. 783

growth and flowering has been the cornerstone of the architec- plant organs but also incorporates a dynamic aspect. It is then
tural school initiated by Hallé and Oldeman during the 70's possible to represent each model by a simplified cartoon
(Hallé and Oldeman, 1970; Hallé et al., 1978). According to the sequence of development (Bell, 1991). Also, as regards the
architectural view, each species, at any given stage of develop- fruit-tree, the observed branch framework of some species may
ment, has a recognizable morphological 'blueprint', or architec- be adequately described by a single model. This is the case for
tural model. Basically, five distinctive morphological and cherry which conforms to Rauh’s model (Lauri, 1992) or apri-
developmental features are used to determine the model of a cot which conforms to Champagnat’s model (Costes, 1993). For
given tree: other species, the architecture is consistent with a combination
− Growth: monopodial or sympodial, i.e. the branch results of two or more architectural models. This is the case for apple
from the development of one meristem (e.g. cherry) or more which combines, to different degrees, depending on the cultivar,
than one meristem (e.g. apricot), respectively. Rauh’s and Scarrone’s features (Lauri and Térouanne, 1995;
− Growth: rhythmic or continuous. This feature is relevant Fig. 1).
only in the Humid Tropics, or in a controlled-atmosphere green- During the course of growth, as the tree matures - with the
house, where growth may potentially occur all year round. transition to the flowering stage - and increases in size, only a
− Branching: rhythmic, continuous or diffuse. This character- part of the buds develop to produce the branching sequence
istic is generally related to growth rhythmicity. consistent with the architectural model. Dormant buds on the
− Growth: orthotropic or plagiotropic, i.e. the branch is
directed vertically upwards or in any other direction, respec-
tively.
− Flowering: terminal (e.g. apple) or lateral (e.g. apple,
peach) on long shoots.
Twenty-three architectural models have been identified to
encompass the hundreds of plant species - trees, shrubs and
herbs - which have been described in such a way. Each model
is labeled with the name of the appropriate scientist. The archi-
tectural model not only provides the spatial arrangement of

Fig. 1. Natural tree habit of a 5-year-old tree of ‘Starkrimson’ on


M7 (A) with lateral and terminal flowering (B). Architectural fea-
tures of apple tree combine architectural models of Rauh and
Scarrone (Photos, credit: PÉ Lauri - INRA). Main architectural
features of the apple tree: 1 - All branches initially orthotropic
(upright trunk and branches). Secondary bending usually occurs Fig. 2. ‘Braeburn’ (A) and ‘Fuji’ (B). Proportions of vegetative lat-
with fruit load. 2 - Branching: rhythmic (A, circle). 3 - Flower- erals (V) and flowering laterals which did not set fruit (F) or
ing: lateral and terminal (B); monopodial/sympodial trunk and which set fruit (A) on 1-, 2- and 3- year-old wood, “1YW”,
branches. NB: Features 1 and 2 are common to Rauh’s and Scar- “2YW” and “3YW”, respectively, on successive annual incre-
rone’s models. Flowering is lateral in Rauh’s model and terminal ments of unpruned fruiting branches (unpublished data from
in Scarrone’s model. Meaux, 1997).
784 Pierre-Éric Lauri

trunk and branches remain dormant. Their development may be and geometry of forest and fruit trees (Costes et al., 1999;
induced in case the existing framework is damaged by pruning, Costes and Guédon, 2002; Godin and Caraglio, 1998; Godin et
bent or favourable growing conditions are created. In either al., 1999) or, more specifically on fruit trees, to analyze shoot
case, depending on its position within the branch framework, architecture (Lauri and Térouanne, 1998), vegetative growth-
the new branching system more or less mirrors the initial archi- fruiting relationships (Lauri et al., 1995, 1996, 1997a, 1997c)
tectural model of the plant. This phenomenon, first described and reactions to bending (Lauri and Lespinasse, 2001).
and termed Reiteration by Oldeman (1974), has been shown to Apple fruiting type - tree ideotype. Independently of the
be a fundamental process by which the plant increases in height architectural analysis of the plant, an apple tree typology was
and volume (Hallé et al., 1978; Bell, 1991). From a practical established by INRA researchers at the end of the 50’s (Bernhard,
point of view, a shoot complex stemming from a pruning cut 1961). Lespinasse and colleagues further extended this work
or a bent shoot is usually considered as a reiterated complex (Lespinasse, 1977, 1980; Lespinasse and Delort, 1986).
(Fig. 3). A large range of cultivars were categorized according to over-
The architectural analysis, i.e. the determination of the archi- all tree habit, i.e. growth direction of branches from upright to
tectural model and reiterated complexes, is the qualitative and weeping, and fruiting type, i.e. position of fruiting on the
first level of architectural investigation. For two decades, a great branch from spur-type (type I) to tip-bearing (type IV) cultivars.
amount of studies have been developed using the methodology
of architecture as a tool to determine morphological categories
(e.g. branching order, growth unit, morphological gradients
along branches) with the objective of modeling tree topology

Fig. 5. Natural extinction (black arrow) and return bloom (“Y”: cur-
rent year flower cluster borne by a bourse and a bourse-shoot in
Year “Y-1”) in Chantecler (credit: PÉ Lauri - INRA).
Fig. 3. The reiteration phenomenon. Reiterated complexes on the
trunk and branches (arrows) in 10-year-old ‘Rome Beauty’ tree
grafted on M7 (credit: PÉ Lauri - INRA).

Fig. 6. Autonomy of the fruiting shoot in Granny Smith. “Y” refers


to the flower-cluster produced the current year. For previous
Fig. 4. Terminal flowering (thick arrows) on upright vigorous years (“Y-1”, and older) only bourse - bourse-shoot complexes
shoots of ‘Granny Smith’ (credit: PÉ Lauri - INRA). are visible (credit: PÉ Lauri - INRA).
J. Kor. Soc. Hort. Sci. 43(6):782−788. 2002. 785

Type I cultivars were characterized by a disjunction between


vegetative growth (upright and vigorous shoots) and fruiting
(fruiting on lateral spurs). Type IV cultivars did not exhibit such
a disjunction and fruiting typically occurred on a large propor-
tion of the growing points, including in terminal position on
long vigorous shoots.
Fruiting type was related to regularity of bearing and as a
general trend, type I cultivars (‘Oregon Spur’, columnar strains)
had a biennial bearing behavior whereas type IV cultivars
(‘Granny Smith’, ‘Red Winter’) were characterized by regular
bearing. This work was further extended by INRA geneticists
and type IV was defined as a low-input regular bearing tree
ideotype suited to facilitate access to the fruit and to reduce
labor costs (Lespinasse, 1992).
An architectural approach to the apple growth and fruiting
strategies. Investigations were carried out on fruiting branches Fig. 7. Variation of the bourse-over-bourse (Number of fruit in Year
of cultivars ranging from type I to IV, to further study how Y which produce a fruiting shoot in Year Y+1/Number of fruit
fruiting developed in one-year-old wood upwards and also to in Year Y) for the successive couples of years of the fruiting
shoot complex and for various cultivars (adapted from Lauri et
better analyze the relationships between growth, branching and
al., 1997a).
fruiting over several years.
Four main features were found to discriminate between cul-
tivars:
− Flowering to vegetative ratio on one-year-old wood of the
fruiting branch. This feature was not related to the above-men-
tioned typology. It had important consequences on the fruiting
behavior of the branch. A high flowering to vegetative ratio was
shown to trigger off an alternation cycle on ‘Braeburn’ whereas
a low flowering to vegetative ratio was related to progressive
start of production for ‘Fuji’ (Fig. 2). This ratio also had con-
sequences on the reaction to bending: bending significantly
increased flowering on one-year-old wood in the case of a high
flowering ratio whereas it had no effect in the case of a low Fig. 8. Theoretical strategies of alternation and non-alternation of
ratio (Lauri and Lespinasse, 2001). fruiting at fruiting shoot and branch levels. Example of 4-year
sequences. A: inflorescence which sets fruit. V: vegetative shoot
− Fruiting in terminal position on vigorous shoots. Type I and (adapted from Lauri and Lespinasse, 1999b).
II cultivars were usually strongly vegetative on upright shoots
whereas type IV cultivars were able to fruit in terminal position were able to string bourse-over-bourse. From a physiological
(Fig. 4). point of view, this autonomy of the fruiting shoot has been asso-
− Bourse-over-bourse, i.e. the ability to fruit in terminal posi- ciated with morphological characters such as the length of the
tion on a bourse-shoot. Average values of bourse-over-bourse fruiting shoot and the volume of the bourse (Lespinasse and
clearly differed depending on the cultivar: type I and II cultivars Delort, 1993; Fig. 6). The reverse situation, i.e. low bourse-
had low bourse-over-bourse whereas type IV cultivars were over-bourse and low extinction, was observed for cultivars such
characterized by a higher value. However, regardless of the cul- as ‘Oregon Spur’.
tivar, bourse-over-bourse generally increased with time, i.e. as In the case of low bourse-over-bourse, two cases were
the fruiting shoot complex became older (Lauri et al., 1997a; observed. In the first case, fruiting of the shoots was synchro-
Fig. 7). nized, leading to the alternating pattern of fruiting. In the sec-
− Extinction, i.e. the physiological abortion of laterals, usually ond case, fruiting of shoots was desynchronized so that a certain
between one- and two-year-old wood (Lauri et al., 1995, 1997c) proportion of shoots produced a fruit in year Y, while the others
(Fig. 5). Type I and II cultivars exhibited low extinction produced a fruit in year Y+1. In this strategy, the relative pro-
whereas type IV cultivars were characterized by a higher value. portions of shoots in the vegetative and fruiting stages, in a
Bourse-over-bourse and extinction thus appeared positively given year, determine the degree of alternation in that year (Fig. 8).
related and generally speaking, cultivars with high bourse-over-
bourse, such as ‘Granny Smith’, were also characterized by A Better Knowledge of Tree Biology to
high extinction (Lauri et al., 1997c; Lauri and Lespinasse, Optimize Tree Management
1999a). At the level of the whole fruiting branch, this strategy
was related to a non-alternating pattern of fruiting: all shoots Traditionally, training system procedures are split into two
786 Pierre-Éric Lauri

successive stages, the establishment of a framework (giving the


tree a given shape: V-Trellis, Vertical Axis, Vase-shape) and the
control of fruiting. The procedures applied at each stage are
based on tree manipulation which now follow well-documented
architectural rules. The building of the trunk and scaffold
branches of a Vase-shape tree typically uses strong reiterated
complexes resulting from winter heading cuts. As long as such
pruning is carried out, the tree is maintained at a stage of vig-
orous vegetative growth which delays the first fruit production.
On the contrary a fast start of production requires minimal
pruning. Therefore, we have to clearly separate actions which
aim at building a given structure, from manipulations which
aim at regulating fruit quantity and quality. In the case where
a self-supporting system is required, the establishment of a
strong structural framework will be done at the expense of fruit-
ing during the first years in the orchard.
In the context of a fast start of production, we consider that
it is important not to strongly modify tree structure in order to
reach the fruiting period as quickly as possible. Once this tree
structure is established, the training procedures have to maintain
an optimal relationship between vegetative growth and fruiting.
Fig. 9. Silhouette of a four-year-old fruiting branch, from ‘Reine
Over the last four decades, INRA proposals for apple tree des Reinettes’ (a), ‘Golden Delicious’-‘Gala’ (b), ‘Granny Smith’
training have evolved from the Vertical Axis (Lespinasse, 1977, (c) and ‘Rome Beauty’ (d). Thick arrows indicate vegetative
1980; Lespinasse and Delort, 1986) towards the Solaxe system growth distribution (adapted from Lauri and Lespinasse, 2000).
(Lespinasse, 1996) and now the Centrifugal training system
(Lauri et al., 2002). At each stage of this progression, training
concepts directly stemmed from better insights into tree physi-
ology. Two examples are proposed to illustrate the direct rela-
tionship between field observations and architectural analysis,
and the change in training proposals.
− The first example is the transition from the Vertical Axis to
the Solaxe systems. The former system was characterized by the
predominance of a single trunk instead of several, as in a tra-
ditional Vase-shape system, and by renewal pruning of the fruit-
ing branch. The latter recommendation involved the pruning of
two-thirds or three-quarters of the three- to four-year-old fruit-
ing branch. This was carried out to avoid the deleterious effects
on fruit of the competition with the vigorous reiterated complex
which usually developed proximally on the upper side of the
initial fruiting branch. Lately it appeared (Lauri and Lespinasse,
2000) that such competition for nutrients and light mainly char- Fig. 10. Tree management in the LITE planting system. Centrifugal
training concept and light distribution in the tree (adapted from
acterized Type I and II cultivars (spur-type cultivars) and to a
Lauri et al., 2002).
lesser extent Type III (‘Golden Delicious’, ‘Gala’; Fig. 9a & b).
For these trees, renewal pruning was aimed at maintaining fruit pendant laterals per se but rather the fact that pendant laterals
in a well-balanced zone with good light distribution and mod- were often in disadvantaged positions underneath upright vig-
erate shoot growth. However, for type IV cultivars with terminal orous shoots. This finding was the basis of a radically new, and
bearing habit, reiterated complexes, when they developed, were reverse concept from that on which renewal pruning was based.
able to fruit in terminal position. As a consequence, the fruit- We proposed to preserve the already established fruiting branch
induced natural bending of all shoots was observed within two and to thin out all reiterated complexes which might develop
to four years after the establishment of the shoot complex, and on it. This free-growing fruiting branch concept (Lauri and
fruit were more equally distributed within the branch volume Lespinasse, 2000) was integrated into the Solaxe training sys-
with less competition between vegetative and fruiting shoots tem.
(Fig. 9c & d). Therefore, from an architectural point of view, − The second example is the already mentioned relationship
we concluded that the relevant characteristic to be controlled by between bourse-over-bourse and extinction which was at the
training was not the presence of fruit on upright, horizontal or origin of the progression from the Solaxe to the Centrifugal
J. Kor. Soc. Hort. Sci. 43(6):782−788. 2002. 787

training systems. In the Solaxe system, fruit-load was adjusted pared to Solaxe system, the Centrifugal training procedure
by classical chemical- and hand-thinning. In the Centrifugal increased light interception by the overall canopy and especially
training system (Fig. 10), fruit-load is controlled by the thin- by fruiting shoots (Willaume, 2002).
ning-out of a certain proportion of fruiting shoots. This proce- The development of the LITE planting system in France
dure, which mimics the natural extinction phenomenon emphasizes the following two aspects, the first of which is the
observed in cultivars such as ‘Granny Smith’ and ‘Red Winter’, greater volume the tree develops with Centrifugal training. We
is called artificial extinction or, in short form, extinction. As a estimate that Centrifugal training increases canopy radius by an
general trend, this procedure improves growth and return-bloom average of 20% as compared to the Solaxe system (unpublished
on the remaining fruiting shoots (Lauri et al., 2002). In this data on 5-year-old ‘Galaxy’ trees; CEHM experimental station,
case, flower- and fruitlet-thinning is used only as a complemen- France). As a consequence, the LITE planting system is prob-
tary tool to control fruit load. When carried out more specifi- ably evolving towards lower density orchards, with larger dis-
cally around the trunk and on the underside of branches as tances between the trees, especially within the row. This aspect,
recommended in Centrifugal training, extinction also improves directly related to light climate, depends on several factors: cul-
light climate within the tree canopy. tivar/rootstock combination, tree height (a high tree may accom-
modate for shorter distances between trees) etc. It is one of our
A Vision for Orchard Management applied research objectives for the years to come.
in the Future ? A second aspect lies more specifically in the concept of tree
training. As evidenced by the name used, traditional systems
The development of sustainable horticulture (in its broad usually refer to a shape which has to be maintained throughout
sense; Tilman et al., 2002) has now become the objective of the entire orchard life-span, i.e. V- or Y-trellis, Vase-shape, Cen-
fundamental and applied research. In the apple tree, it raises the tral Axis, Spindle. The Centrifugal training system refers more
need to better define the type of tree required for a low-input, to a procedure which may therefore be adapted to various tree
regular bearing sustainable apple orchard with high fruit quality. shapes unless incompatibility is observed between procedures
The corpus of knowledge developed on apple tree biology now (extinction may increase vigor when used in combination with
provides some accurate morphological features (e.g. extinction heading cuts). Our main objective is now to improve procedures
and bourse-over-bourse frequency, ability to fruit on vigorous (e.g. bending, extinction) to manage a canopy volume with high
shoots) which may be searched for in progenies along with porosity to light and containing a high proportion of autono-
other functional characteristics such as self-thinning flower-clus- mous fruiting shoots, i.e. with high return-bloom potential
ters, and classical fruit quality and resistance to disease (Lespinasse and Lauri, 1999). In this context, the good man-
attributes (Lauri et al., 1997b). agement of vegetative growth and the progressive start of pro-
The sustainability of orchard planting systems, i.e. growing duction during the first years in the orchard, is of major
high quality fruit on regular-bearing, less-labour demanding importance. Dwarfing rootstocks, such as those belonging to the
trees and with less needs for support system, has also to be M9 type, often lead to an imbalance favouring early excessive
addressed. The improvement of fruit quality is directly related flowering to the detriment of vegetative growth. To overcome
to the light climate within the canopy. A relatively porous can- this problem we are now experimenting Centrifugal training
opy (Lakso, 1994) is required for an optimized balance between concepts on larger self-supporting trees, by using more vigorous
yield and quality. It is still, besides the control of the relation- rootstocks or own-rooted trees (Lespinasse et al., 2000).
ships between vegetative growth and fruiting, an important
objective of all training systems. Shadowing in the interior of Acknowledgements: I would like to thank colleagues from
the canopy makes insect and disease control difficult and the apple MAFCOT (Maîtrise de la Fructification - Concepts et
decreases fruit colour (Dozier et al., 1982), especially in large Techniques) group for helpful discussions and experiments to
canopies. However, training methods which aim at opening tree assess Centrifugal training. I appreciated the comments and
canopy to light, e.g. V- or Y-shaped canopies, generally tend to suggestions by Prof. Yong-Koo Kim of the Department of Hor-
stimulate vegetative re-growth in the tree framework (Lakso et ticulture, Kyung Hee University, Suwon, Korea.
al., 1999). In these cases, global light interception is increased
but on “wrong organs”, i.e. well-illuminated one-year-old vig- Literature Cited
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