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HORTSCIENCE 41(6):1389–1394. 2006.
Graft Compatibility Between Peach
Cultivars and Prunus Rootstocks
Olfa Zarrouk,
1
Yolanda Gogorcena, and Maria Angeles Moreno
Department of Pomology, Estacion Experimental de Aula Dei (CSIC),
Apartado 202, 50080 Zaragoza, Spain
Jorge Pinochet
Agromillora Catalana S.A., C/ El Rebato s/n, 08739 Subirats (Barcelona), Spain
Additional index words. nectarine, SPAD value, stem circumference, plum rootstocks,
incompatibility
Abstract. Trials were established at Aula Dei Experimental Station (EEAD-CSIC,
Zaragoza, Spain) to assess graft compatibility between peach cultivars [Prunus persica
(L.) Batsch] and new Prunus spp. rootstocks or selections. Peach cvs. ÔCatherinaÕand
ÔTebanaÕand nectarine cvs. ÔBig TopÕand ÔSummergrandÕwere grafted on peach
seedlings, plum rootstocks, almond xpeach hybrids, and other interspecific rootstocks.
Part of the evaluated material belongs to the EEAD-CSIC selection program, which has
showed good adaptation to Mediterranean growing conditions. Other rootstocks such as
Bruce, Evrica, Hiawatha, Ishtara, Tetra, and Krymsk-1 have been recently introduced in
Spain. A peach and a plum source, GF 677 and Adesoto 101, respectively, were used as
compatible reference rootstocks. Both are widely used for peach and nectarine pro-
duction in the Mediterranean area.
Most almond xpeach hybrids and slow-
growing plums (i.e., P. domestica and
P. insititia plums like ÔPollizo de MurciaÕ)
were graft-compatible with all tested culti-
vars. However, in the case of fast-growing
plums (P. cerasifera and interspecific hybrids
with this species), performance differed sub-
stantially depending on the evaluated geno-
type. Several levels of response to graft
incompatibility were found for both ‘‘local-
ized’’ and ‘‘translocated’’ types of incompat-
ibility, and some physiological aspects of
graft incompatibility are discussed.
Commercial peach trees [Prunus persica
(L.) Batsch] are usually composed of two
genetically different parts: a scion and a root-
stock. The availability of peach rootstocks
largely depends on the various species or
interspecific hybrids that can be used with
peach as a scion. In the Mediterranean area
(representing 35% of the peach world pro-
duction; FAOSTAT, 2006), almond x peach
hybrids rootstocks are widely used because
of some desirable characteristics such as
tolerance to drought and lime-induced Fe
chlorosis (Socias i Company et al., 1995).
Nevertheless, the highly successful almond
xpeach hybrid rootstock GF 677 is also
extremely vigorous (Wertheim and Webster,
2005; Zarrouk et al., 2005) and relatively
susceptible to nematodes, compact soils, and
waterlogging (Go´mez Aparisi et al., 2001;
Okie, 1987). Because control of tree vigor is
becoming increasingly important for peach
production, plum rootstocks and inter- or
intraspecific plum hybrid rootstocks are used
with peach cultivars. Indeed, plum rootstocks
are generally less vigorous, more tolerant to
waterlogging (Nasr et al., 1977), resistant to
root-knot nematodes (Moreno et al., 1995a;
1995b; Pinochet et al., 1999), and also pro-
vide the possibility to overcome replanting
problems (Nicotra and Moser, 1997) as com-
pared with almond xpeach hybrid rootstocks.
However, the limiting factor for the wide-
spread use of some Prunus spp. for peach
production is the lack of commercial root-
stocks having a wide range of compatibility
with various cultivars (Okie, 1987). For
a composite fruit tree to remain healthy, the
rootstock and the scion should intimately
unite, providing a viable system for the
uptake and translocation of water, minerals,
assimilates, and hormones throughout the
entire lifespan of the plant (Wertheim and
Webster, 2005). Graft incompatibility leads
to poor health, breakage at the graft union,
and premature death or failure of the graft
combination to form a strong and lasting
functional union.
The mechanisms by which incompatibil-
ity is caused and expressed remain unclear
and several hypotheses have been made (Pina
and Errea, 2005). Conversely, previous stud-
ies (Mosse, 1962) described ‘‘translocated’’
graft incompatibility on peach when it was
grafted on several plum rootstocks. Incom-
patibility is usually expressed during the first
year of scion growth in the form of tree
growth cessation and premature defoliation
with leaf discoloration (yellowing or bronz-
ing) (Herrero, 1951). ‘‘Translocated’’ incom-
patibility in peach/plum combinations was
associated with both functional and biochem-
ical alterations at the graft interface (Moing
and Carde, 1988; Moing et al., 1987), in-
ducing a carbohydrate blockage in the scion
above the graft union (Breen, 1975; Moing
et al., 1987; Moing and Gaudille`re, 1992).
Nevertheless, incompatibility symptoms may
occur at a later stage of development (Moreno
et al., 1993), and the presence of some bio-
chemical alterations across the graft union of
Prunus may lead to a slight and delayed
incompatibility as has been described in
cherry by Treutter and Feucht (1991). More-
over, peach/plum combinations can exhibit
symptoms of ‘‘localized’’ incompatibility
(Salesses and Bonnet, 1992). The occurrence
of ‘‘localized’’ incompatibility is character-
ized by anatomic irregularities at the union
interface (Moreno et al., 1995a) with breaks
in cambial and vascular continuity patterns
(Mosse, 1962) and poor vascular connections
(Errea, 2001) inducing mechanical weakness
of the union, which may break after some
years (Herrero, 1951), subsequently leading
to major economic losses.
These problems make rootstock selection
difficult, because commercialization of new
rootstocks requires preliminary evaluation of
possible incompatibility reactions. Addition-
ally, incompatibility can be positively corre-
latedwithwarmclimatesbyincreasingthe
activity of some biochemical substances re-
lated to graft incompatibility(Gur et al., 1968).
This might result in exacerbated graft incom-
patibility when some rootstocks selected in
cold areas are used in warm climate regions.
The objective of this study was to test the
compatibility behavior of several Prunus
rootstocks with peach and nectarine scions
as a preliminary step to their transfer to
commercial peach production orchards.
A rootstock screening experiment was
carried out to identify and determine the graft
compatibility of Prunus rootstocks in the
process of selection and to establish compar-
isons in terms of compatibility with new com-
mercial rootstocks of European, American,
and Russian origins, recently introduced into
the European market. The graft compatibility
of peach (cvs. Catherina and Tebana) and
nectarine (cvs. Big Top and Summergrand)
scions with 44 different Prunus rootstocks
was assessed in nurseries and orchards of
Aula Dei Experimental Station (EEAD-
CSIC). Similarly, some physiological aspects
of incompatibility expression were studied to
search for indicators associated with graft
incompatibility.
Materials and Methods
Plant material
A 3.5-year graft incompatibility study
was carried out at the Aula Dei Experimental
Received for publication 23 May 2006. Accepted
for publication 24 June 2006. Financial support
was provided by Comisio´ n Interministerial de
Ciencia y Tecnologia (AGL2002-4219 and
AGL2005-05533 projects) and by a fellowship
granted to O. Zarrouk from the Agencia Espan
˜ola
de Cooperacio´ n Internacional (AECI). We grate-
fully acknowledge Victoria Fernandez for revising the
manuscript and J. Aparicio and P. Sanchez of the
department of Pomology for their assistance in orchard
management.
1
To whom reprint requests should be addressed;
e-mail z.olfa@eead.csic.es.
HORTSCIENCE VOL. 41(6) OCTOBER 2006 1389
Station. Trials were established on a calcare-
ous soil containing 29% to 30% total calcium
carbonate, 7.4% to 7.6% active lime, and
water pH 8.0 with a clay-loam texture. Peach
and nectarine cultivars were T-budded in situ
in summer of each year from 2000 to 2002.
ÔBig TopÕnectarine, ÔCatherinaÕand ÔTe-
banaÕpeach cultivars were grafted on almond
xpeach hybrids and ÔPollizo de MurciaÕplum
rootstocks. ÔBig TopÕwas also grafted on
interspecific hybrid plums. ÔSummergrandÕ
nectarine was used as an indicator cultivar
for restrictive compatibility (Moreno et al.,
1993), and it was grafted on most rootstocks
in this study. In all trials, the almond xpeach
hybrid rootstock GF 677 was used as refer-
ence because it is commonly used in Medi-
terranean countries, and it is graft-compatible
with all peach cultivars. Some plum root-
stocks such as Adesoto 101 (Moreno et al.,
1995b), Damas GF 1869, and Marianna 2624
were also used for comparison purposes.
The different rootstock species used in
this investigation were obtained from the
rootstock selection program of the Aula Dei
Experimental Station and from Agromillora
Catalana S.A. nursery (Barcelona, Spain).
For practical purposes, rootstock genotypes
were divided into four groups as shown in
Table 1.
Each scion/rootstock combination was
replicated 15 to 30 times depending on the
availability of plant material. Some combi-
nations suffered losses after 3 years of field
testing, mainly as a result of the occurrence of
incompatibilities. Ten replicates per combi-
nation were considered the minimum accept-
able for assessment.
‘‘Translocated’’ incompatibility study
The level of compatibility–incompatibil-
ity was determined during the first 2 years
after grafting by visual diagnosis of the
possible causes of the ‘‘translocated’’ type
of incompatibility in the nursery, e.g., leaf
and wood yellowing and reddening, defolia-
tion, tree vigor reduction, and death (Moreno
et al., 1993). Moreover, a determination of
leaf chlorophyll concentration using a SPAD
502 m (Minolta Co., Osaka, Japan) was made
each year on 1-year-old trees from the end of
June to the beginning of July. This procedure
was used as a potential tool to estimate the
rate of ‘‘translocated’’ graft incompatibility.
Measurements were made on fully expanded
leaves of 10 trees per combination selected
from the middle of the cultivar shoot.
‘‘Localized’’ incompatibility study
When trees were still alive, in the second
and third year after grafting, anatomic exam-
ination of unions (‘‘localized’’ incompatibil-
ity) was carried out. Graft unions were sawed
by a radial–longitudinal plane according to
Mosse and Herrero (1951). The visual rating
of ‘‘localized’’ graft incompatibility was
classified as follows:
Category A = Perfect unions. The line of
union in bark and wood was hardly visible.
Category B = Good unions. The bark and
wood were continuous although the line of
union in the wood was often clearly distin-
guished by excessive ray formation.
Category C = Unions with discontinuities
in the bark. The bark tissues of rootstock and
scion were separated by a dark brown layer of
corky appearance.
Category D = Unions showing vascular
and wood discontinuities. The woody tissues
of rootstock and scion were separated in
many places by clusters of living, nonligni-
fied parenchyma. Bark tissues were generally
as category C.
Category E = Observed breakage of the
tree at the graft union in the nursery.
Also, at the time of internal examination,
stem circumferences 5 cm above and below
the graft union were measured. This method
enabled searching for correlations between
growth characteristics and compatibility–in-
compatibility symptoms.
Analysis of data
Data were evaluated by analysis of vari-
ance with SPSS 13.0 (SPSS, Chicago). Anal-
ysis of variance was made by analysis of
variance at P#0.05 and was used to assess
the significance of stem circumference and
SPAD values. Mean separation was deter-
mined by Duncan’s test and results shown
correspond to mean values. To establish
correlations between incompatibility symp-
toms and stem circumference, the following
scale was designed: level 0 to compatible
grafts, 1 to the presence of only one in-
compatibility type, and 2 to the coexistence
of both incompatibilities.
Results and Discussion
‘‘Translocated’’ incompatibility
As expected, all peach and nectarine trees
on Euamygdalus subgenus rootstocks (Table
1) showed good graft compatibility (Table 2).
Similarly, and with the exception of PP-1 and
PAC 952, most graft combinations were
compatible when peach cultivars were
grafted on slow-growing plums (Table 1).
This was the case of peach and nectarine
cultivars used in this study when they were
grafted on ÔPollizo de MurciaÕplums cur-
rently under selection (e.g., PM 44 AD, PM
95 AD, PM 101 AD, PM 105 AD, PM 137
AD, and PM 150 AD) and on Adesoto 101
used as a reference. Additionally, no incom-
patibility symptoms were observed in Big
Top/St Julien GF 655–2 and in ÔSummer-
grandÕ/Tetra combinations. Results concern-
ing the latter combination are in agreement
with results of previous studies, which re-
ported good compatibility of Tetra with
peach and nectarine cultivars (Nicotra and
Moser, 1997).
In the fast-growing plum group (Table 1),
only three Myrobalan clone rootstocks
(P 2175, P 2980, and P 3293) exhibited good
compatibility when they were grafted with
ÔSummergrandÕnectarine (Table 2). This is in
agreement with the findings of Salesses and
Bonnet (1992) in which Myrobalan P 2175
was tested with other nectarines. The good
compatibility of nectarine cultivars with
some Myrobalan rootstocks support the need
to investigate them with other nectarine and
peach cultivars as a result of their high
resistance and tolerance to some biotic and
abiotic stresses as compared with other plum
rootstocks (Crossa Raynaud and Audergon,
1987). In the interspecific plum group, only
Hiawatha, Ishtara, Jaspi, PAC 941, and PAC
959 showed good graft compatibility with
ÔSummergrandÕnectarine. Similar good com-
patibility behavior results have been previ-
ously observed with Ishtara (Reighard et al.,
1997), Jaspi (Iglesias et al., 2004), and
Hiawatha (Weibel et al., 2003) despite its
parental P. besseyi background, which is
generally graft-incompatible with peach cul-
tivars (Layne, 1987). However, when nectar-
ine cultivars were grafted on fast-growing
plums and interspecific hybrids plums,
‘‘translocated’’ incompatibility increased.
Thus, after the first season of nursery growth,
all combinations of ÔSummergrandÕnectarine
grafted on PP-1, Marianna 2624, Marianna
4001, Myrobalan 29 C, Myrobalan P 1079,
Bruce, Damas GF 1869, Evrica, Krymsk-1,
and Myrobalan GF 3–1 rootstocks (Table 2)
showed clear symptoms of ‘‘translocated’’
incompatibility. The visual symptoms ap-
peared during early and midsummer in the
form of leaf yellowing, a reduction of growth,
and premature defoliation. Cases of incom-
patibility with Evrica rootstock were predict-
able because two of its parents (P. besseyi
and P. cerasifera) are usually known to be
incompatible with peach and nectarine culti-
vars (Layne, 1987). Nevertheless, the incom-
patibility found in Krymsk-1 contrasts with
previous studies carried out in South Carolina
(Reighard et al., 2005). This may be the result
of the differential behavior of this rootstock
depending on pedologic environments and
climatic conditions. We also observed the
development of ‘‘translocated’’ or ‘‘localized’’
incompatibilities when Krymsk-1 rootstock
was grafted with most of the 29 cultivars tested
in another study (data not shown). This sug-
gested that care should be taken in using this
rootstock with commercial peach varieties in
the Mediterranean area.
On the other hand, the severity of in-
compatibility symptoms differed between the
various combinations. ÔSummergrandÕnec-
tarine trees grafted on PP-1, Marianna 4001,
Myrobalan 29 C, and Evrica had a healthy
external bark appearance at the graft union
and homogeneous vigor despite the light
visual ‘‘translocated’’ incompatibility symp-
toms observed in the foliage. In this case, tree
growth cessation was less acute and SPAD
values were not significantly different from
those of compatible trees (Fig. 1). Con-
versely, ÔSummergrandÕtrees grafted on Ma-
rianna 2624, Myrobalan GF 3–1, Myrobalan
P 1079, Damas GF 1869, and Miral showed
premature defoliation, early growth cessa-
tion, very low SPAD values (Fig. 1), and
acute leaf curl since the very first growing
season (1-year-old trees). SPAD values are
generally correlated with leaf chlorophyll
concentration (Shi and Byrne, 1995). Its use
to quantify the rate of leaf yellowing resulting
1390 HORTSCIENCE VOL. 41(6) OCTOBER 2006
from ‘‘translocated’’ incompatibility can be
useful, because low SPAD values may be
associated with the blockage of carbohydrate
assimilation and nitrogen uptake. As the rate
of shoot growth of incompatible graft de-
clines, carbon export from the scion through
the phloem to the rootstock has been reported
to slow down and decrease nitrogen assimi-
lation (Moing and Gaudille`re, 1992; Moreno
et al., 1994). This suggests that the rate of
tissue dysfunctions (Moing and Carde, 1988)
and the degree of leaf chlorosis may differ
from one incompatible combination to an-
other. This different degree of graft incom-
patibility was previously observed in peach
grafted on different Myrobalan clones (Mor-
eno et al., 1993, Yamaguchi et al., 2004) and
may be the result of the differential sensitiv-
ity of rootstocks to poisoning substances
synthesized in peach or nectarine foliage
(Moing et al., 1987). The absence of in-
compatibility in the ÔBig TopÕ/Damas GF
1869 combination (Table 2) contrasted with
previous studies reporting severe incompati-
bility between nectarine cultivars and this
rootstock (Moing and Salesses, 1988). This
may be explained by the different level of
toxic substance synthesis in peach and nec-
tarine cultivars (Moing et al., 1987).
‘‘Localized’’ incompatibility
Like in the ÔtranslocatedÕincompatibility
study, all peach and nectarine trees grafted on
Euamygdalus subgenus rootstocks showed
good graft compatibility (Table 2). Neverthe-
less, in the case of the ÔSummergrandÕ/PAC
960 combination, some gum exudation at the
graft union occurred. The reason for such
exudation remains unknown; however, in
sweet cherry grafts, gum exudation can in-
dicate incompatibility problems (Ja¨nes and
Pae, 2004). Anatomic evaluation of graft
unions indicates ‘‘localized’’ incompatibility
in some 2- to 3-year-old combinations with
several slow-growing plum rootstocks. Graft
unions with ÔCatherinaÕand ÔTebanaÕcultivars
on PM 140 AD (100% for ÔTebanaÕ) rootstock
were classified as ‘‘C’’ (Table 2), which may
be considered the threshold for compatibility
in practical terms. Nevertheless, trees classi-
fied within the ‘‘C’’ category can progress to
an eventual ‘‘localized’’ incompatibility
(‘‘D’’ category) in the future (unpublished
data). Therefore, this material should either
be eliminated from the rootstock selection
process for peach cultivars or be evaluated
for several more years before acceptance.
‘‘Localized’’ incompatibility symptoms were
expressed both in the form of necrosis and
absence of lignified tissues in the wood graft
plane and, in some cases, by the swelling of
the graft union. This was the case of ÔCath-
erinaÕ,ÔTebanaÕ, and ÔSummergrandÕcultivars
grafted on PM 95 AD (Table 2). These cases
of incompatibility with ÔPollizo de MurciaÕ
rootstocks are uncommon. Nevertheless,
because PM 95 AD and PM 140 AD are
open-pollinated selections, they may have an
incompatible parent, which could explain the
results found in this study. On the other hand,
‘‘localized’’ incompatibility was also expres-
sed by union breakage of some 2-year-old
Table 1. Rootstocks used for the peach graft compatibility study.
Rootstock
z
Species Origin
Euamygdalus subgenus
Adafuel P. dulcis xP. persica CSIC, Spain
Adarcias P. dulcis xP. persica CSIC, Spain
GF 677 P. dulcis xP. persica INRA, France
HxM4 P. dulcis xP. persica AC, Spain
Hansen 2168 P. dulcis xP. persica UC, USA
Hansen 536 P. dulcis xP. persica UC, USA
PAC 960, PAC 9501, PAC 9917–01 P. dulcis xP. persica AC, Spain
Barrier P. persica xP. davidiana CNR, Italy
Cadaman Avimag
y
P. persica xP. davidiana INRA, France
Benasque P. persica CSIC, Spain
Missour P. persica Unkown, Morocco
Slow-growing plums
Adesoto 101
y
P. insititia CSIC, Spain
Pollizo de Murcia: PM 44 AD, PM 95 AD,
PM 101 AD, PM 105 AD,
PM 137 AD, PM 140 AD, PM 150 AD
P. insititia CSIC, Spain
PAC 952 P. insititia ? AC, Spain
PP-1 P. domestica ? AC, Spain
St Julien GF 655–2 P. insititia INRA, France
Tetra P. domestica ISF, Italy
Fast-growing plums
Marianna 2624 P. cerasifera xP. munsoniana UC, USA
Marianna 4001 P. cerasifera xP. munsoniana UC, USA
Myrobalan 29 C P. cerasifera GB, USA
Myrobalan P 1079, Myrobalan P 2980,
Myrobalan P 3293
P. cerasifera INRA, France
Myrobalan P 2175 P. cerasifera Unknown, Romania
Interspecific hybrid plums
Bruce P. salicina xP. angustifolia Texas A&M, USA
Damas GF 1869 P. domestica xP. spinosa INRA, France
Evrica (P. besseyi xP. salicina)xP. cerasifera KEBS, Russia
Hiawatha P. besseyi xP. salicina USDA, USA
Ishtara Ferciana (P. cerasifera xP. salicina)x(P. domestica xP. persica) INRA, France
Jaspi Fereley
y
(P. salicina xP. cerasifera)xP. spinosa INRA, France
Krymsk-1
x
P. tomentosa xP. cerasifera KEBS, Russia
Miral P. dulcis xP. cerasifera CSIC, Spain
Myrobalan GF 3–1 P. cerasifera xP. salicina INRA, France
PAC 941 P. dulcis xP. cerasifera AC, Spain
PAC 959 P. domestica xP. insititia AC, Spain
z
Next the rootstock.
y
Protected grant by Community Plant Variety Office (CPVO).
x
Submitted to protection in CPVO.
AC, Agromillora Catalana S.A., private nursery, Spain; CNR, Centro Nacionale della Recerca; CSIC, Consejo Superior de Investigaciones Cientificas; INRA,
Institut Nacional de la Recherche Agronomique; GB, Gregory Brother’s, Calif.; ISF, Instituto Sperimentale per la Fruticultura; UC, Univ. of California; Texas
A&M, Univ. of Texas, College Station; KEBS, Krymsk Experimental Breeding Station. USDA, U.S. Dept. of Agriculture, Mandan, N.Dak.
HORTSCIENCE VOL. 41(6) OCTOBER 2006 1391
Table 2. Graft compatibility and internal examination of the graft unions between peach and nectarine cultivars and Prunus rootstocks.
Cultivar Rootstock
‘‘Translocated’’ incompatibility
symptoms
‘‘Localized’’ incompatibility category
z
ABCDE
Number of trees
Peach
Catherina Adafuel N 20 — — — —
Adarcias N 20 — — — —
GF 677 N 20 — — — —
Adesoto 101 N 30 — — — —
PM 95 AD N — — — 15 —
PM 101 AD N 20 — — — —
PM 105 AD N 20 — — — —
PM 137 AD N 20 — — — —
PM 140 AD N 16 — 4 — —
PM 150 AD N 20 — — — —
Damas GF 1869 N 30 — — — —
Tebana Adafuel N 20 — — — —
Adarcias N 20 — — — —
GF 677 N 20 — — — —
Hansen 2168 N 20 — — — —
Hansen 536 N 20 — — — —
Cadaman Avimag N 20 — — — —
PM 44 AD N 10 — — — —
PM 95 AD N — — — 20 —
PM 137 AD N 10 — — — —
PM 140 AD N — — 10 — —
PM 150 AD N 17 — 3 — —
Nectarine
Big Top Adafuel N 30 — — — —
Adarcias N 15 — — — —
GF 677 N 20 — — — —
Hansen 2186 N 10 — — — —
Hansen 536 N 10 — — — —
Missour N 20 — — — —
Adesoto 101 N 20 — — — —
PM 95 AD N 10 — — — —
PM 101 AD N 10 — — — —
PM 105 AD N 20 — — — —
PM 137 AD N 20 — — — —
PM 140 AD N 10 — — — —
PM 150 AD N 15 — — — —
St Julien GF 655–2 N 10 — — — —
Damas GF 1869 N 10 — — — —
Evrica Ab 15 — 5 — —
Summergrand Adafuel N 30 — — — —
Adarcias N 20 — — — —
PAC 960 N 20 — — — —
HxM4 N 10————
PAC 9501 N 20 — — — —
PAC 9917–01 N 20 — — — —
Barrier N 20 — — — —
Benasque N 10 — — — —
Missour N 20 — — — —
PM 95 AD N — — — 10 —
PM 105 AD N 20 — — — —
PM 137 AD N 10 — — — —
PAC 952 Ab — — — — 15
PP-1 Ab — — — 19 1
Tetra N 10 — — — —
Marianna 2624 Ab — — — 20 —
Marianna 4001 Ab — — — 20 —
Myrobalan 29 C Ab — — — 10 —
Myrobalan P 1079 Ab — — — 10 —
Myrobalan P 2175 N — 10 — — —
Myrobalan P 2980 N 10 — — — —
Myrobalan P 3293 N 10 — — — —
Bruce Ab — — — — 20
Damas GF 1869 Ab 15 — — — —
Evrica Ab 15 — 5 — —
Hiawatha N 20 — — — —
Ishtara Ferciana N 20 — — — —
Jaspi Fereley N — 20 — — —
Krymsk-1 Ab — — — 20 —
Miral Ab 20 — — — —
Myrobalan GF 3–1 Ab 10 — 10 — —
PAC 941 N 10 — — — —
PAC 959 N 10 — — — —
z
Categories A, B, C, D, and E: classification of the rating of ‘‘localized’’ graft incompatibility according to Mosse and Herrero (1951).
N, visual normal trees; Ab, abnormal scion behavior, leaf yellowing, reduction in vigor.
1392 HORTSCIENCE VOL. 41(6) OCTOBER 2006
ÔSummergrandÕnectarine trees when they
were grafted on PAC 952 and PP-1 (Table 2).
The stem diameter growth study (Table 3)
indicates that ‘‘localized’’ incompatibility
was not associated with a decrease in vegeta-
tive growth when dwarfing rootstocks were
used. In fact, 2-year-old trees on Ishtara and
Jaspi showed the lowest circumference below
and above the graft union, but did not signif-
icantly differ from incompatible rootstocks
like Marianna 4001, Bruce, and Krymsk-1.
The same occurred in 3-year-old trees with
Ishtara, which did not differ from trees grafted
on Evrica rootstock. Belonging to the inter-
specific hybrid plum group, Ishtara and Jaspi
rootstocks were compatible when they were
grafted with ÔSummergrandÕnectarine (Table
2). This confirms previous investigations with
other nectarines cultivars (Iglesias et al.,
2004). However, despite having a homoge-
neous appearance, ÔSummergrandÕ/Jaspi and
ÔSummergrandÕ/Ishtara trees were stunted
(Table 3) as compared with other trees with
compatible unions. These results support the
potential use of Ishtara and Jaspi as dwarfing
rootstocks for peach and nectarine cultivars
(Loreti and Massai, 2002; Reighard et al.,
2004). Nevertheless, results in terms of com-
patibility of Jaspi contrast with the report of
De Salvador et al. (2002) in which an in-
compatible behavior of Jaspi with ÔSuncrestÕ
peach cultivar was observed. This suggests
that this rootstock should be tested for longer
time to assess its compatibility behavior with
peach and nectarine cultivars.
‘‘Translocated’’ and ‘‘localized’’
incompatibilities relationship
Some combinations showed the coexis-
tence of two types of incompatibility (Table
2) as reported previously (Moreno et al.,
1995a; Salesses and Bonnet, 1992). This has
been observed in ÔSummergrandÕnectarine
combinations grafted on PAC 952, PP-1,
Marianna 4001, Myrobalan 29 C, Myrobalan
P 1079, Bruce, and Krymsk-1 (Table 2). These
graft unions were classified as ‘‘D’’ and even
‘‘E’’ (smoothly broken unions) with severe
bark anomalies and vascular discontinuities in
the graft plane. ÔSummergrandÕ/PP-1 combi-
nations showed additionally weak swollen and
broken unions. Concerning the group of slow-
growing plums, graft incompatibility was only
found with PP-1 and PAC 952 (Tables 1 and
2). It could be that they hybridized with other
plum species that were incompatible with
peach and nectarine cultivars.
In this study, it was observed that in the
case of coexistence of both incompatibilities,
the ‘‘translocated’’ type preceded the occur-
rence of ‘‘localized’’ incompatibility. This
may confirm that in peach/plum combina-
tions, ‘‘localized’’ incompatibility could be
the result of physiological anomalies at the
graft union incited by ‘‘translocated’’ incom-
patibility. In fact, starch blockage above the
graft union in the scion of incompatible grafts
with ‘‘translocated’’ symptoms (Breen, 1975;
Moing et al., 1987) may prevent cambium
division (Oribe et al., 2003) at the graft
interface and thereby impede vascular tissue
development and successful connection. This
may lead to the formation of discontinuities
in the graft union interface (unpublished
data).
AccordingtoWertheimandWebster
(2005), the trunk diameter above the graft
union of most incompatible combinations is
smaller than below it (Table 3). However,
this difference was not significant in the
present study. Nevertheless, a significant cor-
relation was found between stem circumfer-
ence above the graft union of 2-year-old (r=
–0.524, P#0.01) and 3-year-old trees (r=
–0.238, P#0.05) and both graft incompat-
ibility types, which is in agreement with the
results of Simard and Olivier (1999) for
Fig. 1. SPAD values of ÔSummergrandÕnectarine cultivar grafted on different plum-based rootstocks. Mean separation within columns at P#0.05.
T
‘‘Translocated’’ incompatibility symptoms: abnormal scion behavior, leaf yellowing, and reduction in vigor.
Table 3. Stem circumference (mm) above and below (5 cm) the graft union in ÔSummergrandÕnectarine
grafted on plum rootstocks.
2-year-old tree 3-year-old tree
Rootstock Above graft Below graft Above graft Below graft
PM 95 AD 33.5 b
L
34.3 b
L
——
PM 105 AD 32.0 b 32.3 b — —
PM 137 AD 36.1 bc 39.1 bc — —
PP-1 33.3 b
TL
39.2 bc
TL
——
St Julien GF 655–2 40.6 bc 49.2 c — —
Tetra 39.0 bc 45.8 c — —
Marianna 4001 40.4 bc
TL
38.8 bc
TL
——
Myrobalan 29 C — — 20.0 b
T
31.0 b
T
Bruce 36.6 bc
TL
39.6 bc
TL
——
Damas GF 1869 42.6 c
T
39.4 bc
T
——
Evrica — — 25.1 bc
T
35.8 b
T
Hiawatha 44.4 c 48.5 c
Ishtara Ferciana 30.0 b 36.9 bc 32.5 c 39.7 b
Jaspi Fereley 23.8 a 29.0 ab — —
Krymsk-1 21.6 a
TL
25.6 a
TL
——
Miral — — 22.9 b
T
25.5 a
T
Myrobolan GF 3–1 — — 13.5 a
TL
24.2 a
TL
Mean separation within columns by Duncan’s multiple range tests at P#0.05.
T
‘‘Translocated’’ incompatibility symptoms: abnormal scion behavior,leaf yellowing, and reduction in vigor.
L
‘‘Localized’’ incompatibility occurrence: cambial involutionor/and vascular discontinuity at the graft union.
HORTSCIENCE VOL. 41(6) OCTOBER 2006 1393
apricot. This correlation may be explained by
the decrease of water and nutrient supply
from roots as consequence of graft incom-
patibility, which involves the diminution or
cease of vegetative growth of the scion and
the development of the rootstock as an in-
dependent entity.
In summary, no incompatibility was
found on Euamygdalus subgenus rootstocks
with any of the peach varieties used in this
investigation. This study provides evidence
of the potential use of P. insititia species
rootstocks for the peach industry. Results
showed the possible implication of environ-
mental conditions on the development of
graft compatibility–incompatibility. This
suggests the necessity of investigating ge-
netic and environmental interactions in graft
incompatibility phenomena in Prunus genus.
SPAD values were useful to visually assess
the rate of ‘‘translocated’’ graft incompati-
bility only in cases of severe incompatibility
between scion–rootstock components.
It is concluded that further studies con-
cerning the development of optimal scion–
rootstock combinations based on new plant
material, especially plum rootstocks, includ-
ing P. cerasifera and P. besseyi species,
should be conducted before their commercial
release as rootstocks for peach and nectarine
cultivars.
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