New Zealand Journal of Crop and Horticultural Science

ISSN: 0114-0671 (Print) 1175-8783 (Online) Journal homepage: http://www.tandfonline.com/loi/tnzc20

Grapevine propagation: principles and methods

for the production of high-quality grapevine
planting material

H Waite, M Whitelaw-Weckert & P Torley

To cite this article: H Waite, M Whitelaw-Weckert & P Torley (2015) Grapevine

propagation: principles and methods for the production of high-quality grapevine planting
material, New Zealand Journal of Crop and Horticultural Science, 43:2, 144-161, DOI:
10.1080/01140671.2014.978340

To link to this article: http://dx.doi.org/10.1080/01140671.2014.978340

Published online: 21 Nov 2014.

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 New Zealand Journal of Crop and Horticultural Science, 2015
Vol. 43, No. 2, 144161, http://dx.doi.org/10.1080/01140671.2014.978340

REVIEW ARTICLE
Grapevine propagation: principles and methods for the production of high-quality
grapevine planting material
H Waitea,b*, M Whitelaw-Weckerta,c and P Torleya,b
a
National Wine and Grape Industry Centre, Charles Sturt University, Wagga Wagga, Australia; bSchool of
Agricultural and Wine Sciences, Charles Sturt University, Wagga Wagga, Australia; cNew South Wales
Department of Primary Industries, National Wine and Grape Industry Centre, Charles Sturt University,
Wagga Wagga, Australia
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(Received 27 January 2014; accepted 4 September 2014)

Since the worldwide grapevine planting boom in the 1990s, there have been numerous reports of
sporadic young vine failures and early decline of young vineyards. In many cases, the leading causes
of these problems have been traced to defective, but often asymptomatic, propagating and planting
materials infected with trunk disease pathogens, or with other defects that affect vine establishment,
vigour and longevity. Current propagation practices favour cross-contamination by trunk disease
pathogens and impose physiological stress that affects the quality of finished vines. This review
describes the characteristics of high-quality cuttings and practices that will produce a consistent supply
of quality planting material. The barriers to the production of high-quality grapevine propagating and
planting material are also discussed.
Keywords: grapevine propagation; grapevine quality standards; hot water treatment; nursery sanitation;
trunk diseases

Introduction nurseries to increase production, the quality of

In the past 30 years the global wine industry has
undergone a transformation. It has changed from
an industry characterised by relatively small, tradi-
tionally oriented, family-owned enterprises and a
European focus, to a much more cosmopolitan
industry dominated by multinational corporations.
As a result there is now a stronger focus on quality
assurance and consistency (Aylward 2005). More
wine is now produced and consumed in countries
that have little or no history of wine production
and consumption, and grapes are being grown in
some very challenging climates. However, the
vine propagation industry has not experienced
the same degree of change. It remains largely an
industry dominated by small to medium-sized
family businesses and cooperatives, and although
the progress towards modernisation has enabled
planting material is not yet of a consistently high
standard.
Until recently, there have been few compre-
hensive standards or assessment criteria for grape-
vine material and apart from choice of variety and
rootstock, other important attributes of planting
material have not always been taken into account
by nurseries and grape growers. Consequently,
inferior material has sometimes been planted and
there have been numerous reports from around the
world of vine failures and underperforming vine-
yards needing to be replanted within 510 years of
establishment (Smart et al. 2012). Many of the
failed or underperforming vines were found to be
infected with trunk disease pathogens, or to have
other defects that affect establishment, vigour and
longevity (Stamp 2001; Waite et al. 2013a).

*Corresponding author. Email: hwaite@csu.edu.au

2014 The Royal Society of New Zealand

The losses caused by the failure or poor estab-
lishment and decline of newly planted vines have
become a significant but often unrecognised
burden for both the wine industry and the vine
nursery industry (Scheck et al. 1998; Martelli
1999; Morton 2000; Stamp 2001; Komnek &
Holleinov 2003; Waite & Morton 2007; Rego
et al. 2009; Gramaje & Armengol 2011; Borsel-
lino et al. 2012; Whitelaw-Weckert et al. 2013).
However, there is a growing awareness of the
impact of latent trunk diseases and other defects
on young vines (Morton 2012; Smart et al. 2012;
Smart 2013) and the attention of the wine industry
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is now turning more generally to the quality of
planting material. In Australia, this has prompted
the development of a comprehensive Standard for
Grapevine Material, AS 55882013 (Standards
Australia 2013), and it is anticipated that the qual-
ity of both propagating and planting material will
improve significantly as a result.
Defects in young vines originate from a num-
ber of sources and frequently involve interactions
between several factors including cutting quality,
nursery practices and cold storage conditions
(Stamp 2001; Waite & Morton 2007; Gramaje &
Armengol 2011), but fungal trunk diseases (TDs)
transmitted in asymptomatic propagating material
(Fourie & Halleen 2002) are often primary factors in
vine failure (Gramaje & Armengol 2011; Whitelaw-
Weckert et al. 2013).
The list of TDs transmitted in propagation is
extensive and currently includes Phaeomoniella
chlamydospora, Phaeoacremonium spp., Botryo-
sphaeriaceae spp., Cylindrocarpon spp., Ilyonec-
tria spp. and Phomopsis viticola, which frequently
occur as mixed infections (Larignon & Dubos
2000; Armengol et al. 2001; Aroca et al. 2006;
Halleen et al. 2006; Zanzotto et al. 2007; Rego
et al. 2009; rbez-Torres 2011; Billones-Baaijens
et al. 2013; Whitelaw-Weckert et al. 2013). How-
ever, other pathogens including Fomitiporia punc-
tata (Armengol et al. 2001) and Fusarium spp.
(Halleen et al. 2003) have also been associated with
what is now called young vine decline and it is
likely that more pathogens will be identified as
research progresses. Furthermore, stresses imposed
by poor nursery practices, hypoxic cold storage
Grapevine propagation 145
(Gramaje & Armengol 2012) and poor planting
practicesfactors that are damaging even if trunk
diseases are not presentalso appear to exacerbate
the effects of TDs and result in seriously compro-
mised vines that succumb soon after planting in the
vineyard (Stamp 2001).
Grapevines are relatively easy plants to prop-
agate, but it requires considerable skill and organi-
sation to produce the millions of vines of high
quality that are needed around the world every
year for new plantings and replanting diseased or
uneconomic vineyards. Grapevine propagation
techniques include in vitro propagation (Barlass
et al. 1982), softwood cuttings (Warmund et al.
1986) and field grafting of rooted rootstock cut-
tings (Alley 1957). However, the most usual
method of propagation in commercial nurseries is
by bench grafting dormant one-bud Vitis vinifera
cuttings on to 300400 mm dormant hardwood
cuttings of selected rootstocks and propagating
them as one plant (Nicholas et al. 1992).
To meet the increased demand stemming from
the expansion of the wine industry, modern grape-
vine nurseries have evolved to resemble and
function like factories, with operators working at
single tasks on streamlined production lines. How-
ever, the in-house wastage rates of 40%60%
reported by nurseries worldwide are indicative of
the problems faced by the nursery industry.
Despite considerable progress towards modernisa-
tion (Borsellino et al. 2012), the most significant
challenge remains the ability of nurseries to
maintain a consistent supply of vines that are
sound, healthy and uniform. This will require
detailed descriptions of the practices that result in
the production of high-quality cuttings and vines
coupled with the objective and comprehensive
quality standards for cuttings and finished vines.
Although standards now exist in several juris-
dictions including New Zealand (New Zealand
Winegrowers 2011), Europe (European and Medi-
terranean Plant Protection Organization 1998) and
Australia (Standards Australia 2013), they are not
always comprehensive or supported by best prac-
tice documents. Consequently, defective material
sometimes passes inspection. Here we describe
the characteristics of high-quality cuttings and
 146 H Waite et al.
finished vines, identify the factors that impinge on
quality, and outline the practices that can be uti-
lised to produce a consistent supply of quality
planting material that meets the highest standards.
Characteristics of high-quality grapevine
material
Correct identity, high health status, soundness and
physiological competence are the cornerstones of
cutting and vine quality. Quality cuttings that have
the capacity to become quality vines are uniform,
of verifiable variety and clone, free from serious
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viruses and other pests and diseases, and have
been grown and handled in a manner that preserves
their physiological competence (Nicholas et al.
1992). In addition to the above characteristics,
quality vines must have good plant architecture,
be undamaged and free of physical defects, and
have uncontaminated and fully healed graft unions.
Quality nursery vines establish quickly and per-
form to expectations in the vineyard.
Evaluation of grapevine material
The evaluation of cutting and vine quality is norm-
ally based on a visual assessment when the
material is dormant and without leaves. When
dormant, it is very difficult to identify the variety
and clone of cuttings and vines, and the effects of
stress and latent pests and pathogens are not
readily apparent. These factors have far-reaching
effects on quality. Therefore, close scrutiny of
accompanying documents and labels, dissection of
several samples and sometimes laboratory analysis
may be required to properly evaluate the quality
of cuttings and vines (Chien & Golino 2006;
Stamp 2010).
From an evaluation of many apparently sound
and healthy planting materials supplied by a num-
ber of nurseries, Stamp (2001) reported incomplete
graft healing in 13% of plants and weak or sparse
roots in 4% of rootstock rootlings and 9% of
grafted vines. The percentage of weak, undersized
vines and rootstock rootlings was 3% and 8%,
respectively. Overall, 39% of bench grafted vines
and 35% of rootstock rootlings were deemed
defective. The percentage of defective vines iden-
tified in this work was very high, but reports from
other regions indicated that high percentages of
defects in nursery vines were not unusual (Smart
et al. 2012).
Grapevine material that has major defects such
as graft unions less than 80% healed or lesions on
stems and roots is unlikely to perform to expecta-
tions in the nursery and vineyard. Material that has
minor defects such as graft unions that are between
80% and 100% healed, or sparse, unevenly spaced
roots may be acceptable, but the cumulative effect
of several minor defects is an important consid-
eration when evaluating the risk associated with
such material (Stamp 2001).
Factors affecting the quality of cuttings
and vines
Identity of source cuttings and rooted vines
Centuries of careful selection have given us the
classic cultivars of V. vinifera that produce the
premium wines we prize today (This et al. 2006).
Furthermore, since the arrival of phylloxera (Dak-
tulospharia vitifoliae) in the Old World in the
early 1800s, resistant rootstocks have enabled the
wine industry to continue to survive and prosper
(Bisson et al. 2002). Because planting misidenti-
fied vines can have serious economic conse-
quences, germplasm and mother vine collections
have been established in most wine growing
regions (Laucou et al. 2011) including the Founda-
tion Plant Material Service in the USA (Alley &
Golino 2000), the regional Vine Improvement
Schemes and Associations in Australia (Cirami
et al. 1988), ENTAV-INRA in France (Grenan
et al. 2000), the Vine Improvement Association of
South Africa (Fourie & Halleen 2004a) and the
New Zealand Grapevine Improvement Group
(Hayward et al. 1998). These valuable collections
have been established to supply propagators
with certified, true-to-type cuttings that are also
free of serious viruses. They are regularly mon-
itored for incorrectly identified vines and virus
symptoms and the propagating material sourced
from them has been pivotal in reducing the effects

of grapevine viruses and the problems caused by
misidentified vines on the wine industry. How-
ever, certification is not always a guarantee of
physiological competence or that the material is
free from other diseases and physical defects
(Chien & Golino 2006). Furthermore, the use of
certified cuttings for propagation is not universal,
particularly when demand for cuttings outstrips the
supply of certified material (Waite et al. 2013a).
Soundness and physiological competence
Soundness and physiological competence are
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essential characteristics of quality propagating
and planting material. Physical defects, including
wounds caused by wire rub, hailstones, vermin or
machinery, incomplete healing of graft unions,
restrictions caused by tendrils or twine, damaged
or blind buds and frost-damaged tissue all impinge
on the quality of cuttings and the finished vines.
They also cause structural weakness and provide
entry points for pathogens (Stamp 2001). Physio-
logical competence affects the underlying regen-
erative capacity of the material.
Intrinsic physiological characteristics that en-
able a cutting or vine to establish well and grow
vigorously are critically influenced by genotype,
environmental factors (Winkler et al. 1974) and
management, and are established in the previous
season while the cane is growing on the mother vine
(Samish & Spiegel 1957; Nicholas et al. 1992). The
poorer rooting capacity of the rootstocks Ramsay
(Smith et al. 2013) and 140 Ruggeri (Kracke et al.
1981) compared with other rootstock varieties is a
direct function of genotype, but the majority of
factors that affect regenerative capacity are a func-
tion of complex interactions between genotype,
environmental conditions and mother vine manage-
ment (Hartmann et al. 2001).
Inadequate or excessive water and fertiliser, and
temperature extremes in mother vine plantings all
affect the regenerative capacity of cuttings taken
from them (Nicholas et al. 1992). Excessive
applications of nitrogen and water result in rank
cuttings that have elongated internodes and rela-
tively large pith diameters that strike poorly (Pearse
1943). Equally, cuttings from drought-stressed
Grapevine propagation 147
vines or under-fertilised vines that have borne
heavy crops are generally thin and stunted, and
also root poorly (Nicholas et al. 1992; Smart et al.
2002). If they do develop roots, such cuttings are
likely to succumb to stress and make unsatisfactory
growth in the nursery.
Reduced photoperiod during shoot growth,
thought to result in lower carbohydrate stores
from reduced photosynthesis, has also been
reported to have a negative impact on the rooting
of cuttings (Hansen et al. 1978; Hartmann et al.
1990). This factor may also explain the poorer
performance of cuttings from the distal ends of
canes relative to cuttings from the basal ends
(Nicholas et al. 1992), but there is little evidence
for a direct correlation between starch reserves and
rooting of cuttings (Treeby & Considine 1982).
Carbohydrates are primarily stored in the ray
tissue of 1-year-old canes and ripening of the
canes does not occur until all the ray tissue is
packed with starch grains (Mullins et al. 1992).
Consequently, it is rare for a fully ripened dormant
cutting of 7 mm diameter to lack adequate
reserves for rooting and graft healing and it is
possible that other, as yet unidentified physiolo-
gical factors affect the regenerative capacity of
cuttings (Treeby & Considine 1982; Smart
et al. 2002).
The lower strike rates in some rootstock
varieties, particularly Vitis berlandieri hybrids,
have been linked to constitutively high levels of
the plant hormones gibberellic acid (GA) and
abscisic acid (ABA). Soaking cuttings in water
to leach these inhibitors has been a popular remedy
(Kracke et al. 1981), but soaking can compromise
the quality of cuttings by spreading disease (Waite
et al. 2013a) and, if prolonged, flooding the tissue
with water, causing hypoxia and initiating dam-
aging fermentative respiration (Vartapetian &
Jackson 1997). Hormone levels in cuttings change
naturally over the storage period and propagation
is often more successful if cuttings are processed
late in the dormant season rather than in early or
mid-winter (Alley 1979; Nicholas et al. 1992).
In commercial nurseries, where many thousands
of cuttings are propagated over an extended
season, it is not always practical to delay
 148 H Waite et al.
processing. However, rooting can be promoted
without recourse to soaking: for example, by the
application of synthetic auxins in the form of
indolebutyric acid (IBA) to the bases of cuttings
of V. berlandieri hybrid rootstocks which are
difficult to root (Nicholas et al. 1992). The use
of IBA makes little difference to the more easily
rooted rootstock varieties (Nicholas et al. 1992)
and V. vinifera cuttings root so easily that rooting
hormones are rarely used.
Pests and diseases
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The impact of pests and pathogens transmitted in
propagation on the establishment and longevity of
vines is well documented (Morton 2000; Andret-
Link et al. 2004; Retief et al. 2006; Rego et al.
2009), but the concept of mother vine plantings
and vine nurseries as hubs for the dispersal of
other pests and diseases across national and inter-
national boundaries (Martelli 1999; Jeger et al.
2007) is less well recognised.
For viruses, primary control measures have
focused on the production of clean certified
mother vines grown from propagating material
following thermal therapy (Over de Linden &
Chamberlain 1970; Barlass et al. 1982; Golino
2000; Gribaudo et al. 2006). This method has
been used to establish foundation plantings free
of serious viruses (Cirami et al. 1988; Alley &
Golino 2000; Grenan et al. 2000). However,
foundation plantings are vulnerable to infection
in the field (Walter & Martelli 1998). Although
certified mother vines generally have a low titre of
pests and diseases, there is a smaller but real risk
of transmitting latent pests and pathogens such as
phytoplasma diseases, crown gall, Pierces dis-
ease, mildews, botrytis blight, mealy bug, nema-
todes and fungal trunk diseases (Barlass et al.
1982; Rego et al. 2009) in certified propagating
material.
Furthermore, asymptomatic but infected cut-
tings from mother vines infected with TD patho-
gens can act as sources of inoculum during the
propagation process resulting in cross infection of
entire batches of cuttings and the finished nursery
vines grown from them (Fourie & Halleen 2004a;
Billones-Baaijens et al. 2013). TD pathogens have
multiple host species (Crous & Gams 2000;
Halleen et al. 2004; Slippers et al. 2004) and are
spread by rain splash, wind and contaminated soil
(Gubler et al. 2004; Halleen et al. 2007; Ampon-
sah et al. 2009). These diseases are thus less easily
managed in mother vines than viruses that are
transmitted in propagation and by mealy bug and
other controllable coccid vectors (Sforza et al.
2003). Because TDs are spread by wind and rain
splash it is rare to find mother vine plots that
are free from infection and it must therefore be
assumed that all batches of cuttings, including
those from certified sources, carry some TD
inoculum (Fourie & Halleen 2004b).
Symptoms of TDs in nursery vines include
poorly healed graft unions with dark brown or
black staining in the wood extending away from
the wound, and dark staining and streaking in the
woody tissue of the trunk extending away from
disbudding wounds and upwards from the base
of the vine (Stamp 2001; Wallace et al. 2004).
However, small, highly localised areas of dark
tissue around the base of a vine and at the graft
union and disbudding wounds are a normal part of
wound healing and are not usually cause for
concern (Stamp 2001).
Hot water treatment (HWT), usually but not
universally at 50 C for 30 min, is currently the
most effective control for some trunk diseases
including crown gall and nematodes and other
pests and pathogens, in cuttings and finished vines
(Lear & Lider 1959; Ophel et al. 1990; Crous
et al. 2001; Fourie & Halleen 2004a,b; Gramaje
et al. 2009a). However, hot water treated material
is susceptible to stresses caused by inappropriate
handling practices (Gramaje & Armengol 2012),
and because HWT does not always provide 100%
control of TD pathogens (Rooney & Gubler
2001), its use remains controversial (Waite et al.
2013a).
Mother vine management and cutting harvest
The impact of mother vine management and
harvesting practices on the quality and physiolo-
gical competence of cuttings and vines cannot be

underestimated. A poor quality cutting is very
unlikely to become a quality vine. However, little
attention has been paid to the role of mother vine
management in the production of quality propag-
ating material, and there is a paucity of literature
on the subject.
In Australia, rootstock mother vines are not
usually trellised, but allowed to sprawl along the
ground from a self-supporting crown approxi-
mately 30 cm above the soil surface. Although
this method of production is inexpensive and rela-
tively easily managed, it may favour infection by
trunk disease pathogens (Whiteman et al. 2007).
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However, experimental evidence is lacking. Trel-
lising systems for mother vines that position the
crowns and canes well above the soil surface are
used in New Zealand and some European coun-
tries including Germany, Italy and Spain, but they
are not universal. Furthermore, there is little pub-
lished information on the effects of other manage-
ment practices such as irrigation and nutrition
regimes for cutting production. However, fully
lignified and apparently normal cuttings taken
from V. vinifera mother vines that have borne a
very heavy crop often perform poorly and should
be avoided (Hartmann et al. 1990). The reason for
this is not known. It has been speculated that
reduced starch stores in the wood are involved, but
other less obvious and more complex physiological
changes are also likely to be involved, including
the carbonnitrogen balance and its influence on
carbon partitioning between roots, shoots and fruit
and thus on ABA signalling (Morinaga et al. 2003;
Grechi et al. 2007).
Nursery management
High-quality propagating material, appropriate and
timely processing, scrupulous sanitation (Hartmann
et al. 1990) and meticulously kept records are the
cornerstones of successful propagation. Good nur-
sery practices preserve and enhance the quality of
cuttings as they proceed through the propagation
process, but quality propagating material can be
rendered defective by inappropriate or careless pra-
ctices in the nursery (Stamp 2010).
Grapevine propagation 149
The fundamentals of nursery management
have been well known since the 1940s (Baker
1957) and details of best practice can be found in
the many texts that have been published on the
subject (Baker 1957; Hartmann et al. 1990). None-
theless, good nursery practices have not always
been communicated clearly and in a form mean-
ingful for grapevine propagators, and conse-
quently inadvertent cross-contamination and
physiological stress are common features of cur-
rent practice in the vine nursery industry. Grape-
vine nurseries can seriously compromise the
quality of the finished vines with some common
practices including cold storage of wet cuttings
with free water and sawdust or other unsterile
material added to the storage bags and by sealing
bags to completely exclude air, soaking cuttings in
water and erratically applied sanitary practices
(Waite & Morton 2007). These practices are dis-
cussed below.
Sanitation
General sanitation in the nursery is most critical to
the maintenance of quality in propagation (Daugh-
trey & Benson 2005), but often not well un-
derstood. Sanitary standards in the vine nursery
industry have improved, but general sanitary pro-
cedures are not yet applied consistently through-
out the propagation process (Waite et al. 2013a).
Inadvertent contamination of grapevine material
from sources including dust, water, tools, callus-
ing media and clothing is common (Whiteman
et al. 2004; Edwards et al. 2007; Vigues et al.
2009; Aroca et al. 2010; Billones-Baaijens et al.
2013; Cardoso et al. 2013).
Soaking cuttings to compensate for dehydra-
tion that might have occurred during transport and
handling and to leach growth-inhibiting hormones
is widely believed to be beneficial to cuttings and
often used repeatedly during the propagation pro-
cess (Waite et al. 2013a). There is some evidence
to support the use of soaking to promote cutting
establishment (Alley 1979; Gramaje & Armengol
2012), but soaking severely compromises sanita-
tion by providing multiple opportunities for micro-
organisms from the tissue and bark of infected
 150 H Waite et al.
cuttings to contaminate the hydration water and
from there to infect propagation wounds on unin-
fected cuttings (Baker 1957; Whiteman et al.
2004; Gramaje & Armengol 2011; Waite et al.
2013b).
Trunk disease inoculum has been found in
water used for soaking cuttings in several coun-
tries including New Zealand (Whiteman et al.
2004; Billones-Baaijens et al. 2013), Spain (Aroca
et al. 2010), Portugal (Cardoso et al. 2013), South
Africa (Retief et al. 2006), Australia (Edwards
et al. 2007), Italy (Pollastro et al. 2009) and
France (Vigues et al. 2009). Although nurseries
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now generally use potable water and often add
biocides (Waite et al. 2013a), the potential for
cross-contamination remains if the biocide con-
centration is too low, resistant microorganisms are
present (Fourie & Halleen 2004b) or multiple
batches of cuttings are soaked in the same water.
Microorganisms can also form biofilms that per-
sist on the surfaces of soaking vessels after routine
cleaning and potentially contaminate subsequent
batches of cuttings (Costerton et al. 1994; Epstein
et al. 2011; Waite et al. 2013b).
Stressful conditions
Sources of stress in the nursery include dehydra-
tion, exposure to extremes of temperature, soak-
ing, wounding stress, and exposure to toxic fumes
and agrochemicals. These are often overlooked as
dormant cuttings are not actively growing and
thus do not appear to be alive to the nursery
personnel. Furthermore, the effects of stress may
be delayed or compounded (Lichtenthaler 1996),
which makes it difficult to identify the original
causes, particularly if they are viewed as minor
and inconsequential.
Dehydration can occur whenever cuttings are
left exposed to the air, especially during the post-
callusing period when the shoots and leaves are
emerging. However, soaking cuttings to reverse
the effects of dehydration floods the cutting tis-
sue with water which has an inherently lower
oxygen concentration than air, potentially trigger-
ing fermentative respiration if soaking is pro-
longed or the cuttings are prevented from drying
to the normal moisture levels for dormant cuttings,
usually 50% of fresh weight (Lavee & May 1997).
In the vineyard, bud break in vines occurs
before root growth commences (Mullins et al.
1992) and this sequence also occurs in grafted and
ungrafted cuttings. Under normal field conditions,
buds become hypoxic towards the end of the
dormant season, prompting fermentative respira-
tion that stimulates loosening of the bud scales
and bud burst and thus a return to aerobic respira-
tion (Halaly et al. 2008; Ophir et al. 2009). Buds
of cuttings undergo the same sequence of events,
but placing cuttings in sealed bags for long-term
cold storage will promote and prolong hypoxia.
Furthermore, sealed storage bags inhibit the dis-
sipation of ethanol and acetaldehyde, the toxic by-
products of fermentative respiration (Woodstock
& Taylorson 1981; Kimmerer & Kozlowski 1982).
These fermentation products combined with the
oxygen deficit can impair the regenerative capa-
city of cuttings. The woody tissue of cuttings and/
or young vines that have suffered prolonged
hypoxia has been observed to be a characteristic
grey-green colour rather than the true green or
cream of healthy tissue. It has been observed by
the author and remarked on by nursery managers
that cuttings or vines with these symptoms die
soon after planting out, or lack vigour, or suffer
from greatly delayed bud burst and eventually fail.
The high temperature of HWT can also be a
cause of stress. Although research has demon-
strated that HWT has a minimal impact on the
viability of cuttings and nursery vines (Orffer &
Goussard 1980; Wample et al. 1991; Wample
1993; Caudwell et al. 1997; Waite & May 2005;
Gramaje et al. 2009a; Gramaje & Armengol 2012),
there are anecdotal reports of significant losses of
cuttings and finished vines occurring after HWT
in commercial situations. In the relatively cool
climate of New Zealand, unacceptable damage to
vine tissue has been reported after treatment at
50 C for 30 min; treatment at the slightly lower
temperature of 48 C for 30 min has been found to
be safe and effective against some pathogens
(Graham 2007). However, in the relatively hot
climate of Spain, vines and their pathogens are
acclimated to high temperatures and 50 C for

30 min is insufficient to control TD pathogens,
and the higher temperature of 53 C is used
without significant damage to the treated material
(Gramaje et al. 2009a).
There is also evidence that pre- and post-HWT
practices, particularly cold storage, affect the
response of vine material to HWT (Gramaje &
Armengol 2012). The relatively high temperature
of HWT has been reported to damage material that
has not had sufficient time to acclimate to ambient
temperature after removal from cold storage
(Lindquist 1986). Acclimation to ambient allows
the material to express some protective heat shock
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proteins that confer improved tolerance to the
unnaturally high temperature of HWT (Morrell &
Wample 1995). However, anecdotal reports from
industry indicate that cuttings begin to emerge
from dormancy if the acclimation period is
extended beyond 810 h, making them susceptible
to damage from HWT.
Hot water treated cuttings are particularly
susceptible to fermentative respiration in storage
if they are placed in sealed bags within 24 h of
treatment. The respiration rates of hot water
treated cuttings are very high in the immediate
post-HWT period (Waite 2005) and it is probable
that available oxygen in sealed bags is quickly
consumed, creating hypoxic conditions earlier
than normal in the storage period. A sudden return
to aerobic conditions after the opening of storage
bags may also trigger the generation of damaging
reactive oxygen species from electrons that accu-
mulate in the respiratory chain during fermentative
respiration (Vartapetian 2006).
Gramaje and Armengol (2012) found that
long-term storage of cuttings for 4 weeks or more
immediately after HWT was detrimental to cutting
growth and establishment. There are also anec-
dotal observations from nurseries of strong
winey aromas indicative of fermentative respira-
tion when sealed bags of hot water treated cuttings
are opened after long-term storage (6 weeks or
more). However, the effects of cold storage on
cuttings can be variable. Wample (1997) and
Waite and May (2005) reported that when hot
water treated cuttings were able to cool to ambient
and respiration returned to pre-HWT rates before
Grapevine propagation 151
packaging, placing in cold storage for 46 weeks
was not detrimental. In commercial situations
where cuttings are sometimes placed in cold
storage before their respiration rate has returned
to the pre-HWT rate, and cold storage conditions
are not always ideal, treating cuttings post-storage
reduces the risk of fermentation provided that they
are not repackaged and returned to storage.
Stress also favours disease development indir-
ectly (Schoeneweiss 1981). The contribution of
stress from inappropriate handling practices in the
nursery, particularly soaking and anaerobic storage
conditions, or prolonged storage for more than
68 weeks (Probst et al. 2012) may be a factor in
the aggressive development of trunk diseases that
occurs in young vines towards the end of the
growing season. Stressful conditions are known to
affect trunk disease development in the field
(Graniti et al. 2000), but further investigation is
required to determine if stressful conditions during
propagation also affect trunk disease development
in the nursery.
Processes for the production of high-quality
nursery vines
Clearly defined, standardised and consistently
applied standard operating procedures (SOPs) are
the foundation of uniform, high-quality, high-
value vines (Baker 1957), but they are not a
common feature of grapevine nurseries (Waite
et al. 2013a). Although nursery and vine accred-
itation schemes operate in several jurisdictions
(European and Mediterranean Plant Protection
Organization 1998; New Zealand Winegrowers
2011; Standards Australia 2013), they often lack
detail, particularly in the area of general nursery
sanitation and stress management. Consequently,
nursery practices are variable within and between
nurseries (Waite et al. 2013a), wastage rates are
high, and the quality of the resulting planting
material is inconsistent (Stamp 2001; Smart 2013).
Standard operating procedures founded on
best practice are the cornerstones of waste reduc-
tion (Hartmann et al. 1990). In vine nurseries they
can prevent the incursion, establishment and
transmission of pests and diseases (Baker 1957;
 152 H Waite et al.
Daughtrey & Benson 2005). Effective SOPs detail
the practices in every step of the propagation
process from cutting harvest to the sale of the
finished product and identify the personnel
responsible for their implementation, management
and periodic review (Baker 1957). Furthermore,
SOPs should have enough flexibility to allow for
minor modifications that suit the needs of indi-
vidual nurseries.
Cutting harvest and handling
The very first step in the propagation process is
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the harvesting of cuttings and their transport from
the mother vine block to the nursery. Well-
managed harvesting operations in mother vine
blocks are critical to the maintenance of cutting
quality (Daughtrey & Benson 2005). Cuttings left
lying on the vineyard floor are liable to suffer
dehydration, contamination by soil borne organ-
isms and frost damage. Transport to the nursery
may take several days, further increasing the risk
of damage through dehydration, adverse tempera-
tures and hypoxia. If the ambient winter temper-
ature is above 24 C, the respiration rate of the
material increases. As a result, the available oxy-
gen in the packaging is quickly consumed and
metabolic heat or the products of fermentative
respiration are unable to dissipate, which compro-
mises the viability of the cuttings.
Grading and storing cuttings
The most effective method of preventing trans-
mission of pests and diseases in propagation is to
ensure only healthy cuttings enter the nursery
(Daughtrey & Benson 2005), and for this reason
cuttings sourced from a certification scheme are
preferred. If uncertified cuttings are used, it is
advisable for a sample to be tested for the presence
of viruses and trunk disease pathogens by an
accredited laboratory before propagation begins.
In addition, the normal variability within each
batch of cuttings means that grading of each batch
is necessary to remove defective or diseased
cuttings at the beginning of the propagation chain.
This reduces the cost burden associated with
processing cuttings that will not become saleable
plants (Baker 1957). Grading cuttings for size also
increases efficiency by reducing downstream
handling and costs associated with in-batch
variability.
Cuttings that are not to be processed immedi-
ately after grading can be stored in a clean, cool
room at 12 C in clean packaging with several
small, well-spaced, 710 mm holes that allow air
to reach the cuttings without danger of dehydra-
tion. Soaking cuttings in water or packing them
with added water or moisture-retaining materials
such as sawdust promotes the growth of undesir-
able microorganisms, particularly Botrytis cinerea
(Baker 1957). To prevent the growth of moulds on
cuttings in cold storage, cuttings have previously
been soaked in 0.5% Chinosol (8-hydroxyquino-
line sulphate). However, the use of Chinosol is
not absolute proof against the growth of undesir-
able microorganisms. Since it has also been
reported to inhibit callusing and graft healing
(Becker & Hiller 1977), the use of Chinosol
has declined. In some nurseries, the use of Chino-
sol has been replaced by a brief fungicidal dip or
spray applied to cuttings and storage containers.
However, only the organisms susceptible to the
fungicides are inhibited and undesirable mould
growth can occur, particularly if the cutting sur-
faces are wet.
Storage temperatures below 1 C can cause
physiological stress from freezing, and uncon-
trolled temperature fluctuations above 2 C can
have a serious impact on cutting quality (Hartmann
et al. 1990). When storage temperatures exceed
4 C, growth of microorganisms is promoted; the
increased metabolic activity of the cuttings also
generates heat, consumes available oxygen and
leads to fermentative respiration (Becker & Hiller
1977). Other sources of stress in cold storage
include fluctuations in metabolic rate caused by
moving vine material in and out of storage and
exposure to high levels of ethylene when vine
material is stored with climacteric fruit such as
apples and pears (Pierik et al. 2006).

Hot water treatment
Routine hot water treatment of all cuttings prior
to propagation is the most effective means of
controlling some trunk diseases in asymptomatic
material, particularly if used in conjunction with
fungicide drenches (Fourie & Halleen 2004b;
Fourie & Halleen 2006; Gramaje et al. 2009b).
However, HWT is a significant stress and should
only be applied to cuttings of sound and healthy
appearance.
Treatment time and temperature depend on
climate and vary from 48 C for 30 min in cool
climates (Graham 2007) to the standard 50 C for
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30 min in moderate climates (Waite & Morton
2007) and 53 C for 30 min in warm climates
(Gramaje et al. 2009a). In France, 50 C for 45
min is used to ensure that the eggs of phytoplasma
vectors are also destroyed (Caudwell et al. 1997).
HWT is usually followed by an equal cooling time
in clean water at ambient temperature. However,
water cooling is a potential route of recontamina-
tion, particularly if the water has been used mul-
tiple times, is not of potable quality, or has been
exposed to wind- or water-borne contaminants.
Following HWT, cuttings should be allowed to
recover by being loosely covered in a clean and
protected environment for 24 h to become surface
dry before further processing or packaging for
storage. However, current information indicates
that the storage life of hot water treated cuttings
and vines is shorter than that of untreated material
(Gramaje & Armengol 2012). It is therefore pre-
ferable to treat cuttings after storage and immedi-
ately prior to processing, provided that they are
allowed to come to ambient temperature for 812 h
before treatment.
There is no standard design for HWT plants,
which have various sizes and heating methods.
HWT plants and equipment for commercial situa-
tions are designed to function effectively and
efficiently. Even heating of water that is circulated
by means of a pump ensures that it returns to the
specified temperature less than 2 min after the
baskets of cuttings are inserted. Tank capacity is
usually in the region of 30005000 L, to accom-
modate 500010,000 cuttings, allowing about
Grapevine propagation 153
0.5 L per cutting. Hot water tanks, cool down
tanks and dipping baskets are constructed of
stainless or galvanised steel to prevent corrosion,
and hot water tanks are insulated to prevent heat
loss. Prior to treatment, accurately calibrated
temperature probes are inserted in a minimum of
three zones (top, middle and bottom) and con-
nected to data loggers that record the temperature,
date, time and duration of each treatment. To
improve efficiency, dipping baskets are designed
to sit away from the bottom and sides of the tank
and are packed so that the cuttings or vines lie in
the same direction as the flow of water circulated
by the pump. Very tight packing of cuttings or
vines has been found to impede water flow and
heat transfer.
Grafting and callusing operations
Grafting and the preparation of cuttings for
callusing are critical stages in the propagation
process and necessitate making wounds that are
inherently vulnerable to biotic and abiotic con-
tamination unless very stringent sanitary standards
are maintained (Baker 1957; Fourie & Halleen
2004b). Contaminated wounds and poorly matched
graft unions fail to heal properly, remain open to
infection and create structural weaknesses in the
finished vines (Stamp 2001). General sanitary
measures, including regular and thorough cleaning
and tidying of nursery facilities and equipment,
appropriate and timely disposal of plant waste and
rubbish and managing waste water and runoff
(Baker 1957; Stewart-Wade 2011) are essential
elements of pest and disease control in vine nurs-
eries. Designated grafting rooms that are not open
to rain and wind-blown dust, or birds and other
vermin, reduce the risk of contamination. Con-
sumables should be kept in a designated store-
room away from vine material (Baker 1957).
During propagation, grafting rooms should be
cleaned frequently. Benches, grafting machines
and tools should be disinfected with appropriate
products such as sodium hypochlorite at every
meal break and at the end of each day. However,
practices such as sweeping or hosing the floor
should only be done at the end of the day when
 154 H Waite et al.
the benches have been cleared as they raise clouds
of dust or water droplets that can contaminate
everything in the grafting room (Baker 1957).
Other apparently inconsequential actions that
compromise the quality of the finished vine and
contribute to losses during propagation include
delays in processing caused by meal breaks and
exposure to high temperatures, agrochemicals, or
ethylene from the exhaust of forklifts (Winkler
et al. 1974; Hartmann et al. 1990; Nicholas et al.
1992; Saltveit 1999; Waite & Morton 2007).
However, holding cuttings and pre-cut scion
buds before grafting in water is the practice most
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likely to result in universal cross-contamination.
Soaking has been shown to significantly increase
the titre of microorganisms in cutting tissues
because fungi and bacteria from the wood and
bark of some cuttings can enter the water and
infect propagation wounds (Waite et al. 2013b).
Instead, cutting scion buds immediately before
grafting, covering rootstock and scion cuttings
with a clean damp cloth, and implementing mea-
sures to avoid delays in processing prevents
dehydration and minimises the risk of cross-
contamination (Hartmann et al. 1990).
For callusing, cuttings are normally packed
into clean boxes containing a moist, but not wet,
sterile medium such as perlite or vermiculite, and
then incubated for 23 weeks at 2629 C until a
ring of callus forms around the base of the cuttings
and the graft unions. Callusing is slower at lower
temperatures (2627 C), but slower growing
callus forms a stronger union than callus that
develops at higher temperatures (2829 C). If
cuttings are left in the callusing boxes for more
than 3 weeks, excessive amounts of callus tissue
can form that impedes the formation of new xylem
and phloem across the graft union. Callus tissue
should not protrude outwards from the graft union
by more than 23 mm (Hartmann et al. 1990).
Callusing cuttings at temperatures above 29 C, or
high-density packing of cuttings, can prevent the
dissipation of metabolic heat with fatal conse-
quences for the cuttings.
The warm, dark and humid environment in
callusing rooms is particularly favourable for the
growth of some pathogens (Hartmann et al. 1990).
Therefore, disinfecting callusing boxes, regular
and thorough cleaning of callusing rooms, and
prompt disposal of spent callusing media are
essential elements in pest and disease control
(Baker 1957). However, sweeping or hosing floors
and walls can spread pathogens in raised dust and
airborne water droplets. Mopping or washing
floors and walls is preferable. Graft unions can
be protected from dehydration during callusing by
wrapping in budding tape or dipping in grafting
wax (Becker & Hiller 1977), but callus develop-
ment and graft healing require high levels of
oxygen, and heavy waxing of the graft union
prior to callusing can impede graft healing (Hart-
mann et al. 1990). Healing is also impeded if wax
penetrates the graft union and forms a barrier
between the rootstock and scion. Tight packing of
cuttings in callusing boxes also prevents air
movement around the cuttings, favours growth of
pathogenic fungi (Hartmann et al. 1990) and
inhibits the dissipation of potentially fatal meta-
bolic heat generated by the cuttings during callus-
ing. Monitoring of temperatures in callusing boxes
is therefore advised.
Post-callusing operations
In any batch of callused cuttings there may be some
that do not meet quality standards. Damaged or
diseased cuttings and cuttings with incomplete graft
unions should be removed to reduce processing
costs and disease pressure (Nicholas et al. 1992).
Callused cuttings require careful handling to
avoid losses or checks to growth caused by trans-
planting shock. Cuttings are vulnerable to dehyd-
ration until roots form and it may be beneficial to
harden (acclimate) the buds of callused cuttings
before they are planted into containers or a field
nursery. Hardening can be done by removing the
top few centimetres of callusing medium to
expose the buds and graft unions and holding the
callusing boxes in a well-lit but protected envir-
onment for 2 or 3 days (Nicholas et al. 1992).
If the graft union and bud have not already
been waxed, the tops of cuttings can be dipped in
a thin layer of grafting wax to prevent dehydration
(Nicholas et al. 1992). Grafting wax also provides

a measure of frost protection if the callused
cuttings are planted directly into a field nursery.
However, excessively thick wax is undesirable
(Hartmann et al. 1990) and treatment with an anti-
transpirant product such as the acrylic polymer
Envy may be preferable to additional layers
of wax.
The quality of potting mixes and field nursery
soils is critical to cutting establishment. Potting
mixes are normally soilless (Baker 1957) and are
rarely sources of weeds or soil-borne pathogens
(Daughtrey & Benson 2005). Supplementary fer-
tilisers with moderate levels of nitrogen and
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moderate amounts of irrigation water applied at
frequent intervals prevent stress and promote
steady, moderate growth (Hartmann et al. 1990).
However, excessive applications of water and
nitrogen promote rank, soft growth.
Field nursery soils should be well drained,
friable and free of serious weeds, nematodes and
pathogens (Hartmann et al. 1990). Transmission of
black foot pathogens (Cylindrocarpon spp. and
Ilyonectria spp.) is a particular problem in field
nursery soils (Whitelaw-Weckert et al. 2013) and
reducing the titre of these pathogens in the soil can
be difficult (Agust-Brisach & Armengol 2013).
However, it may be possible to reduce disease
inoculum in nursery soils by growing a rotation
crop of mustard as a biofumigant (Bleach et al.
2010). It may also be helpful to use a fungicidal
dip just before callused cuttings are planted out to
reduce infection through the small wounds made
when fragile root initials are broken during
planting (Alaniz et al. 2011).
Shoot growth is usually trimmed during the
growing season to facilitate management and
prevent overcrowding which promotes the devel-
opment of fungal diseases. However, a regular
spray programme is also necessary to prevent
diseases such as powdery and downy mildews and
botrytis blight (Hartmann et al. 1990). Pruning the
roots of vines in field nurseries mid-season, by
running a tractor-drawn blade through the soil
approximately 30 cm either side of the vine row,
promotes the formation of a fibrous root ball and
facilitates lifting and handling at the end of the
season, but the process of trimming the roots and
Grapevine propagation 155
shoots when the vines are lifted to facilitate
handling also creates wounds that may be vulner-
able to infection by soil-borne pathogens.
After the vines are dug from field nurseries,
damaged and sub-standard vines should be dis-
carded and their roots and shoots lightly trimmed
to reduce excessively long roots and shoots that
interfere with handling. The roots are particularly
vulnerable to dehydration if left exposed to the air
(Hartmann et al. 1990). However, sitting the roots
in water creates anaerobic conditions in the roots
and promotes the growth and transmission of soil-
borne pathogens. Covering the vines with clean
damp cloths and prompt processing prevents
dehydration and avoids the negative effects of
soaking (Baker 1957).
There are two main methods for storing bare-
rooted vines: cold storage for long-term storage
and heeling in for short-term storage. For heel-
ing in, bundles of trimmed vines are placed in
trenches and the roots loosely covered with damp
uncontaminated earth, sand or sawdust. However,
the shoots of heeled-in vines soon begin to grow,
and cold storage is the preferred method for
holding vines in the dormant state for more than
1 or 2 weeks (Hartmann et al. 1990).
Packaging and storage conditions for vines
correspond to those for cuttings. Vines that are to
be held in cold storage are washed to remove soil
and plant debris, allowed to drain and may be
trimmed again to further reduce the root and shoot
systems for ease of packaging (Hartmann et al.
1990). However, trimming is a stress and causes a
short-term rise in respiration rates (Macnicol 1976)
and oxygen consumption. Therefore, freshly
trimmed vines should be allowed a period of
recovery before packing and placing in cold
storage to prevent the development of fermentat-
ive respiration. Comments from the nursery indus-
try have also indicated that placing untrimmed or
lightly trimmed vines in cold storage, removing
them for trimming before despatch, and then
returning them to storage is detrimental for the
same reasons and is therefore not recommended.
A final grading of finished vines before despatch
and dissection of a representative sample for
signs of physical damage, trunk diseases and
 156 H Waite et al.
contamination in the graft union also helps to
prevent the sale of sub-standard vines (Nicholas
et al. 1992).
Record keeping
Although rarely stated overtly in the literature,
record keeping and decision-making management
tools underpin successful propagation. A well-
designed record keeping and information manage-
ment system that identifies and traces vine mater-
ial through the whole propagation process, from
the mother vines through to the sale of finished
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vines, is an essential tool for managing costs and
detecting actual and potential problems (Fonsah
2006). It is the foundation of quality management
and compliance systems. However, keeping and
managing records can be seen as a tedious task
and for that reason needs to be purposeful, stream-
lined and easy to manage (Rhigetti & Halbleib
2000; Srensen et al. 2011).
There are many record keeping and analysis
systems that employ both paper-based and elec-
tronic techniques, including bar coding systems
(Senneset et al. 2007). Choice of a system depends
on the size and structure of individual businesses,
but regardless of type, record-keeping systems
need to provide useful and readily accessible
information (Rhigetti & Halbleib 2000). Time
devoted to the collection, maintenance and ana-
lysis of appropriate records is repaid many times
over in reduced losses and increased profitability
(Rhigetti & Halbleib 2000; Fonsah 2006).
Discussion
Although data detailing the economic and social
effects of poor planting material on the wine and
propagation industries are limited, the issues
associated with poor planting material are begin-
ning to receive coverage in the wine industry press
(Stamp 2010; Smart 2013) and moves to improve
the quality of planting material are gathering
momentum. However, the low cost of planting
material, the underestimation of its true worth, and
the disjointed and fragmented nature of the vine
propagation industry are universal impediments to
change in an industry that is strongly rooted in
tradition and often escapes the notice of people in
the wine industry education and research sectors.
Given the dual imperatives of economic and
environmental sustainability, the wine industry
now demands a more professional and exacting
approach to the selection and management of
propagating and planting material (Bisson et al.
2002). There is much to be done, both at the local
level and internationally. The comprehensive writ-
ten standards for grapevine material that are
emerging around the world (European and Medi-
terranean Plant Protection Organization 1998;
New Zealand Winegrowers 2011; Standards Aus-
tralia 2013) and the principles and processes
outlined here are the essential first steps on the
road to the propagation of quality planting mater-
ial, but meaningful and sustained change requires
a multifaceted and collaborative approach invol-
ving all sectors of the grape and wine industry.
These include holders of germplasm collections,
suppliers of cuttings and nurseries, as well as
grape growers, researchers, extension staff and
funding bodies. Every district and jurisdiction has
its own approach to the issues associated with sub-
standard planting material, but the multinational
nature of the wine industry mandates a common
approach to the core issues of vine health, identity
and soundness that transcend local and interna-
tional boundaries.
Of the many challenges associated with devel-
oping a unified approach to vine propagation,
quality of cuttings is a particularly difficult issue
because it involves the financing, ownership and
management of germplasm collections and multi-
plication blocks. Many of these are managed by
industry groups and volunteers, and are under
threat as a result of funding shortfalls (McMichael
et al. 2013).
Another important issue that has a great
impact on the quality of grapevine planting
material is nursery sanitation, particularly the
practice of soaking cuttings in water, and hypoxic
cold storage conditions. However, changing
entrenched practices that are regarded as beneficial
(Kracke et al. 1981; Gramaje et al. 2009a) can be
a difficult task (Willock et al. 1999; Piderit 2000),

particularly when the perceived benefits of these
practices such as advanced bud break and the
prevention of dehydration are apparent soon after
treatment, but the detrimental effects of delayed
establishment and increased pathogen load are not
apparent until after the material has left the nursery
(Stamp 2001; Oliveira et al. 2003; Fourie &
Halleen 2004b; Smart 2013; Whitelaw-Weckert
et al. 2013).
The criteria set out here for evaluating the
quality of grapevine propagating and planting
material and the methods for producing it are, on
the face of it, straightforward and evolutionary
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rather than revolutionary, but there are complex
underlying structural issues in the vine propaga-
tion industry and the broader wine industry that
make their implementation far from certain. Build-
ing a sound and practical internationally focused,
industry-wide framework for the production of
high-quality propagating and planting material
will be a slow and painstaking procedure that
will require much discussion, debate, testing and
assessment of the risks and benefits of a more
directed approach to vine propagation. Much
goodwill, courage and a clear vision will be
required to bring about fundamental improve-
ments in planting material, but success will bring
with it a new era in grape and wine production.
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