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mycorrhizal inoculation



 
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https://hal.archives-ouvertes.fr/hal-00885762/document
Micropropagation of walnut trees (Juglans regia L) and
response to arbuscular mycorrhizal inoculation
R Dolcet-Sanjuan, E Claveria, A Camprub´ı, V Esta´un, C Calvet
To cite this version:
R Dolcet-Sanjuan, E Claveria, A Camprub´ı, V Esta´un, C Calvet. Micropropagation of walnut
trees (Juglans regia L) and response to arbuscular mycorrhizal inoculation. Agronomie, EDP
Sciences, 1996, 16 (10), pp.639-645. <hal>
HAL Id: hal-00885762
https://hal.archives-ouvertes.fr/hal-00885762
Submitted on 1 Jan 1996
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agronomie: plant genetics and breeding
Micropropagation of walnut trees (Juglans regia L) and
response to arbuscular mycorrhizal inoculation
R Dolcet-Sanjuan E Claveria A Camprubí V Estaún C Calvet
1 Departament de Genètica Vegetal; 2 Departament de Patologia Vegetal, Institut de Recerca i Tecnologia Agroalimentaries,
Centre de Cabrils, 08348 Cabrils (Barcelona), Spain
(Received 3 July 1996; accepted 23 September 1996)
Summary &mdash; Juglans regia clones were micropropagated through the establishment of in vitro shoot-tip cultures and
induction of adventitious roots. Shoot proliferation of juvenile J regia material was higher than with clones established
from adult trees. Genotype and juvenility of the plant material were determinant factors of the in vitro rooting ability.
Rooting of adult J regia clones was improved after rejuvenation by subsequent subcultures. The sucrose concentration
in the root elongation medium affected the formation of secondary roots per rooted shoot and the acclimatization of
plants. Mycorrhizal inoculation of micropropagated walnut trees improved post-acclimatization growth in greenhouse
conditions. Inoculation with the arbuscular mycorrhizal fungi Glomus mosseae or G intraradices significantly improved
plant survival when transferred to nursery.
walnut / micropropagation / rooting / acclimatization / arbuscular mycorrhiza
Résumé &mdash; Micropropagation du noyer (Juglans regia L) et effet de l’inoculation avec des champignons
mycorhizogènes à arbuscules. Des clones de noyer, Juglans regia, ont été micropropagés via l’établissement de cultures
in vitro d’extrémités de tiges et l’induction de racines adventives. La prolifération des tiges a été plus grande pour
les clones juvéniles que pour les clones établis à partir d’arbres adultes. L’aptitude d’enracinement in vitro a varié avec
le contexte génétique et le stade, juvénile ou adulte, du matériel de base. Le taux d’enracinement de clones adultes de
J regia a augmenté après rajeunissement du matériel au moyen de subcultures successives. La concentration de
sucrose dans le milieu d’élongation des racines a eu un effet sur le nombre de racines formées et sur le sevrage des
plantes. La mycorhization des noyers micropropagés a amélioré la croissance des plantes en serre. L’inoculation avec
les deux champignons mycorhizogènes essayés : Glomus mosseae et Glomus intraradices a augmenté le taux de
survie des noyers lors de leur transplantation en pépinière.
acclimatation / enracinement / micropropagation / mycorhizes à arbuscules / noyer
* Correspondence and reprints
INTRODUCTION
The micropropagation of walnut trees presents some difficulties that have to be solved before
this technology can be used as a commercial tool
for clonal propagation of selected clones
(McGranahan et al, 1987; Rodriguez et al, 1989;
Gruselle and Boxus, 1990). The main difficulties
are: i) the establishment of in vitro shoot-tip cultures
from adult walnut trees, ii) uneven and low
rates of multiplication or rooting and iii) high number
of losses during acclimatization of plantlets to
soil.
Walnut trees (Juglans nigra x J regia) were
first micropropagated through in vitro culture of
zygotic embryos and multiplication by shoot-tip
cultures (Cornu and Jay-Allemand, 1989; JayAllemand
et al, 1992). Explants from vigorously
growing seedlings were used in the micropropagation
of the walnut rootstock Paradox (J hindsii
x J regia) (Driver and Kuniyuki, 1984). However,
little progress has been made in the micropropagation
of adult walnut trees since the methodology
published by McGranahan et al (1988). A
modification of this methodology was used here
to establish shoot-tip cultures and further micropropagate
selected adult J regia clones. The
effects of rejuvenation and sucrose level on the
in vitro rooting ability and acclimatization of J
regia clones were determined.
The use of micropropagation techniques
where plants are grown under axenic conditions
and then transplanted to sterile media, lacking
arbuscular mycorrhizal (AM) fungi, do not allow
the formation of the mycorrhizal symbiosis.
During the first stages of growth the high nutrient
levels of the potting media masks the mycorrhiza
effect in many instances. However, some of the
problems reported after field transplanting, like
stunting and low survival rates, might be due to
the lack of symbiosis. In this study the convenience
of AM inoculation to increase the survival
and growth of J regia plants after transfer to nursery
was assessed.
MATERIALS AND METHODS
Plant material
Mature trees of the Juglans regia clones ’Serr’ and
’MB-T-231’ were selected for their characteristics as
cultivars and rootstocks (Aletá and Ninot, 1993).
Scions were collected and grafted on seedlings of the
same species. Two to 3-year-old grafted trees, maintained
in a greenhouse to reduce the risk of in vitro
contamination, were used as source of explants. Eight
juvenile J regia clones (’SBE4’, ’SBE5’, ’SBE11’,
’SBE15’, ’SBE21’, ’SBE22’, ’SBE26’, ’SBE27’) were
established from embryonic axis. Zygotic embryos
originated from open pollinated crosses with ’MB-T-
231’ as the maternal component.
Medium and culture conditions
After a preliminary comparison between DKW-C medium
(McGranahan et al, 1987), MS medium (Murashige
and Skoog, 1962) and woody plant medium (WPM)
(Lloyd and McCown, 1980), the DKW medium (Driver
and Kuniyuki, 1984) was chosen for the establishment
and multiplication phases. This medium contained the
original mineral nutrients, with 0.29% (w/v) Phytagel,
and a pH of 5.5. Growth regulators (N6-benzyladenine
[BA] and indole-3-butyric acid [IBA]) were added to the
multiplication medium before autoclaving. In the root
induction phase, &alpha;-naphthaleneacetic acid (NAA) was
dissolved in dimethyl sulfoxide (Dolcet-Sanjuan and
Claveria, 1995) and added to the medium after autoclaving.
During establishment, shoot multiplication and
root elongation, all cultures were kept at 28 °C under a
photoperiod of 16 h of cool-white fluorescent light
(70 &mu;mol m-2 s-1). Root induction was performed in
the dark.
Explant establishment
and shoot multiplication
Establishment of in vitro shoot-tip cultures from mature
walnut trees was accomplished following the same
methodology described for Pistacia vera L (DolcetSanjuan
and Claveria, 1995) and walnut (McGranahan
et al, 1988). Each explant was cultured in a tube containing
15 mL of DKW medium with 5 &mu;M BA and
0.05 &mu;M IBA. During the first 8 weeks, shoot-tip
explants were subcultured weekly. Previous to their
mass multiplication, apparently uncontaminated shoots
were screened for bacterial contamination in plated
523 medium (Viss et al, 1991). Explants that showed
bacterial growth on this medium were discarded.
Depending on the genotype, branching and multiplication
started between 3 to 9 months after initiation. For
shoot multiplication, shoots or segments of at least
1 cm long were transferred every 4 weeks to flasks
containing 100 mL of DKW medium supplemented with
5 &mu;M BA and 0.05 &mu;M IBA. Seven segments were
transferred to each flask.
Juvenile J regia clones were established from
embryonic axes isolated and established in vitro as
described by Jay-Allemand and Cornu (1986). After
elimination of bacterial contaminated cultures and taking
into account the in vitro multiplication and the rooting
rates, following the methodology described by JayAllemand
et al (1992), eight clones were selected and
used in rooting and acclimatization trials.
Shoot multiplication experiments
The effects of juvenility and genotype on the multiplication
rates were studied. Shoot-tip cultures from eight
embryo-derived J regia clones and two mature J regia
clones (’Serr’ and ’MB-T-231’) were used. Each experi- ment consisted of a completely randomized design
with ten replicate flasks containing five shoots each
per clone. The multiplication rate was recorded, in ten
consecutive subcultures (fifth to 15th), as number of
shoots longer than 1 cm per cultured shoot after 4
weeks of culture.
Rooting experiments
A sequence of two rooting media, one with auxin and
another with vermiculite (Jay-Allemand et al, 1992), was chosen for rooting assays. The rooting medium
used was DKW (Driver and Kuniyuki, 1984) with onefourth
the concentration of macronutrients and with no
glutamine. Root induction was performed by culturing
the shoots in the dark, for 5 days, in the rooting medium
supplemented with 25 &mu;M IBA. Each flask contained
100 mL of medium and five shoots (3 to 5 cm
long) taken from stock multiplication cultures. Root
elongation was achieved after culture for 3 weeks in
auxin-free medium and normal light conditions. The
root elongation medium contained a 55% (v/v) vermiculite
(250 mL vermiculite and 200 mL modified DKW
medium per flask).
The in vitro rooting ability of eight juvenile clones
(’SBE4’, ’SBE5’, ’SBE11’, ’SBE15’, ’SBE21’, ’SBE22’,
’SBE26’, ’SBE27’) and two mature clones (’Serr’ and
’MB-T-231’) was examined. Depending on the clone,
shoots from the fifth to the 35th multiplication subculture
were utilized. After root elongation the percentage
of shoots forming roots was scored.
The beneficial effects of rejuvenation on rooting were determined by in vitro micrografting the mature
J regia ’Serr’ on seedlings of the same species.
Micrografts were performed with the help of a silicon
tube to hold together scion and rootstock. When the
scion was elongated enough, it was excised, subcultured
and a second grafting was performed. Root
induction was done in microcuttings derived from
none, one or two micrografts. After root elongation the
percentage of shoots forming roots was scored.
The effects of subsequent multiplication subcultures
on rooting of mature (J regia ’Serr’ and ’MB-T-231’) or
juvenile (’SBE11’, ’SBE26’, ’SBE27’) clones were also
studied. The number of subcultures in multiplication
medium depended on the clone and ranged from six to
55. The percentage of shoots forming roots was
scored after root elongation.
The effects of three sucrose levels (30, 15 and 0
g·L) on the root elongation of J regia ’Serr’ were evaluated.
Shoots were grown for 5 days on the auxin-containing
medium described earlier and then on auxinfree
medium for 3 weeks. The percentage of shoots
forming roots and the number of secondary roots per
rooted shoot were scored.
For each experiment, three to five flasks with five
shoots each were randomly selected for each treatment
level. The results were analyzed statistically by
analysis of variance and treatment means were compared
by Fisher’s least significant difference test and
planned contrasts (Petersen, 1985).
Acclimatization experiments
Rooted plantlets were washed and potted in trays with
250 mL connical pots, containing steam-sterilized
(100 °C, 1 h) substrate (2 peat:1 perlite, v/v). Plantlets
were acclimatized for 3 weeks in plastic boxes. Two
fungicide treatments (benomyl or captan) were applied
during the first 2 weeks of acclimatization. They were
gradually exposed to reduced relative humidity by progressively
removing the box covers during the last
week.
The effect of the sucrose level (30, 15 and 0 g·L)
during the in vitro root elongation on the number of
acclimatized J regia ’Serr’ plants was determined.
Shoots were grown on auxin-containing medium for 5
days and then on auxin-free medium for 3 weeks. Four
weeks after the acclimatization phase, the number of
plants that developed new leaves and a root system was recorded and the percentage of acclimatization
was calculated.
Inoculation with AM fungi
Four weeks after acclimatization, J regia ’Serr’ plants
with new leaves and an extensive root system were
repotted into 4 L polypropylene containers with a peat
and sand mixture (2:1, v/v) and were inoculated with
AM fungi. Inocula consisted of mixed soil and mycorrhizal
roots of Allium porrum L. Plants colonized by
Glomus mosseae (BEG12) (Nicol and Gerd)
Gerdemann and Trappe or Glomus intraradices
(BEG72) Schenck and Smith were grown on sandy
soil. Each plant was inoculated with 10 g of the appropriate
inoculum placed under the root system at the
moment of planting. A filtrate of soil inoculum free from
AM fungi was added to the control treatment. There
were 30 plants per treatment. After 6 months’ growth in
a greenhouse, the plants were transferred to the nursery.
The percentage of plant survival was assessed
after 3 months field growth. Root samples were collected
and stained with 0.05% trypan blue in lactic acid
(Koske and Gemma, 1989), and the percentage of AM
root colonization was determined using the grid-line
intersect method (Giovannetti and Mosse, 1980). The
percentage of plant survival after the transplant to
nursery soil conditions was assessed after 3 months’
growth.
RESULTS AND DISCUSSION
Explant establishment
and shoot multiplication
To reduce the risk of in vitro contamination, grafted
trees maintained in the greenhouse were the
best source of material for the establishment of
shoot-tip cultures from field-grown, adult, selected
trees. Most shoot-tip cultures established from
cuttings or directly from field-grown trees were
ruined by internal contamination of explants.
Secretion of phenolics during the first days of culture
was solved with frequent subcultures into
fresh medium. In spite of this problem, establishment
and multiplication of mature J regia ’Serr’
and ’MB-T-231’ were successfully accomplished
with the methodology described here. Juvenile J
regia clones, established from embryonic axes,
presented less problems of contamination or
secretion of phenolics during establishment than
mature plant material. In addition, multiplication
rates of mature J regia shoot-tip cultures were
satisfactory but never as high as those of some
juvenile J regia clones (fig 1). Significant differences
on multiplication rates were also found
among juvenile J regia clones. Morphological differences
such as apical dominance, axillary
branching and leaf size were observed among
embryo-derived J regia clones. On the basis of
these reported differences, three clones
(’SBE11’, ’SBE26’ and ’SBE27’) were selected
and micropropagated for latter agronomic evaluation.
Adventitious root formation
The use of a vermiculite containing medium during
the root elongation phase improved rooting of
J regia clones (Jay-Allemand et al, 1992). The
number of roots, root elongation and formation of
secondary roots was improved by the addition of
vermiculite to the gelified medium. In these conditions,
the formation of callus in the shoot base
was reduced and a good vascular connection
with the root system was found (Heloir et al,
1994).
Significant differences in root formation were
observed among embryo-derived J regia clones.
Three of them, ’SBE11’, ’SBE26’ and ’SBE27’,
reached values from 25 to 40% of shoots forming
roots. Higher rooting percentages were obtained
with J nigra x J regia clones (Jay-Allemand et al,
1992) than with embryo-derived J regia clones. In
early stages of the multiplication phase (fifth to
35th subcultures), the percentage of shoots forming
roots was lower for mature J regia ’Serr’ and
’MB-T-231’ (12 and 5%, respectively) than juvenile
J regia clones (fig 2). Large variations in the
rooting aptitude were observed between successive
subcultures. Selection of microcuttings during
sequential rooting assays could be an explanation
for these differences.
Embryo-derived clones showed good
response to root induction even at early stages of
multiplication. The number of subcultures in the
multiplication medium progressively rejuvenated
the plant material and also increased the rooting
aptitude of the mature clones ’Serr’ and ’MB-T-
231’ (fig 3). However, more than 35 subcultures
were necessary before J regia ’Serr’ reached a
40% rooting, similar to that obtained with J regia
embryo-derived clones only after five subcultures
in the multiplication medium. With the objective of
accelerating the rejuvenation process, J regia
’Serr’ derived from adult material was micrografted
on shoots of embryo-derived cultures. The
micrografting technique was successful; however,
the percentage of rooting was not significantly
increased after one or two successive micrografts
(20 and 29%, respectively) when compared
with the rooting of no micrografted material
(34% of shoots forming roots).
The percentage of microcuttings forming roots
as well as the number of primary roots per rooted
shoot increased when the sucrose level in the
root elongation medium was reduced from 30 to
15 g·L (Dolcet-Sanjuan et al, 1997). The beneficial
effects of a lower sucrose level (15 g·L) were
also found in the number of secondary roots per
rooted shoot of J regia ’Serr’ (fig 4A). This characteristic
was associated with a better acclimatization
of plantlets to soil conditions when 0 or 15
g·L sucrose were added (fig 4B). However, 0 g·L sucrose had a lower percentage of rooting than
15 g·L (Dolcet-Sanjuan et al, 1997).
Acclimatization of plantlets
The percentage of acclimatized plants was genotype-dependent,
J regia clones being more easily
adapted to soil conditions than J nigra x J regia
hybrid clones (Frossard et al, 1997).
Acclimatization assays were done with more than
800 J regia plantlets, which resulted in the survival
of 25% of ’MB-T-231’ and 83% of ’Serr’ (out
of 16 and 359 plantlets, respectively). Regardless
of the genotype, the addition of vermiculite to the
root elongation medium (Jay-Allemand et al,
1992; Heloir et al, 1994) favored the formation of
primary and secondary roots. Moreover, lowering
the sucrose content in the root elongation medium
presumably stimulated the photosynthetic
capacity, and consequently the survival of
plantlets during acclimatization to soil was
improved (fig 4). Acclimatized J regia plantlets
grown for over 1 month in a 2 peat: 1 perlite substrate
produced some new leaves and an extensive
root system, adequate for inoculation with
AM fungi.
AM fungi on plant survival
Early AM inoculation of micropropagated J regia
’Serr’ trees improved the post-acclimatization
growth, in greenhouse conditions. Inoculation of
J regia ’Serr’ with the AM fungi Glomus mosseae
or G intraradices significantly improved plant survival
when transplanted to the field. Only 20% of
the control plants survived while 60 and 80% of
plants colonized with G mosseae and G
intraradices, respectively, came through the
transplant.
AM root colonization of micropropagated J
regia ’Serr’ trees (fig 5) was 72 ± 15% for G
mosseae, 73 ± 11 % for G intraradices and 51 ±
5% for uninoculated plants. Control treatments
were colonized naturally after field transplanting,
but the benefits of the colonization were
observed only for the plants previously inoculated
and transplanted with the symbiosis.
The results indicate that although AM fungi are
present in the soil and the roots are eventually
colonized with native endophytes, the early inoculation
of micropropagated walnut plants with
selected AM fungi confers protection against the
stress situation due to the transplant process.
The absence of AM fungi in the roots of micropropagated
plantlets affects both survival and
growth of micropropagated J regia trees following
their transfer to field conditions. Survival increase
can justify the inoculation of in vitro propagated
walnut plants with AM fungi in the early phases of
plant growth after acclimatization.
ACKNOWLEDGMENTS
This work was supported by a grant from INIA
(Instituto Nacional de Investigaciones Agrarias) and
the AIR Program of the European Commission (CT92-
0142). Use of trade names does not imply endorsement
of the products named nor criticism of similar
ones not named.
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Dolcet-Sanjuan R, Gruselle R, Jay-Allemand C, MeierDinkel
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Dolcet-Sanjuan R, Claveria E (1995) Improved shoottip
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Driver JA, Kuniyuki AH (1984) In vitro propagation of
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27P, 42
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Dünya florasının yaklaşık %10’nunda oluşan ektomikoriza başlıca Pinaceae
(çam, göknar, karaçam, ladin baldıran), Fagaceae (meşe, kestane, kayın),
Betulaceae (kızılağaç, huş) Salicaceae (kavak, söğüt), Juglandeceae (Amerikan
cevizi, pekan), Myrotaceae (okaliptüs) ve diğer bazı ağaçlarda görülür. Birçok
mantar ektomikoriza oluşturucu olarak tanımlanmıştır. Bu mantarlardan çoğunun
üreme yapıları veya meyve evleri olarak şapkalı mantarlar, puf topları, veya truffle
üretirler. Bunlar meyve evlerinde rüzgarla, yağmurla ve memelilerle uzun
mesafelere taşınabilen sporlar üretirler. Ektomikorizal kolonizasyon emici köklerin
şeklini ve rengini değiştirir. Onlar çatallanarak dallara ayrılabilir, çok dallı veya
diğer şekillerde olabilirler. Onların rengi siyah, kırmızı, sarı, kahverengi, beyaz
veya bu renklerin karışımları olabilir (Marx 2001).
Başa dön
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Yeni Konu Gönder   Cevap Gönder    Mesaj Panosu -> Ceviz Genel Saatler GMT +1 zaman dilimine göredir
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Etiketler : mycorrhizal inoculation

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