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Selection and nursery production of

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Article in Journal of Horticultural Science and Biotechnology · January 2006

DOI: 10.1080/14620316.2006.11512022

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Journal of Horticultural Science & Biotechnology (2006) 81 (1) 3–17

Review Article

Selection and nursery production of ornamental plants for

landscaping and xerogardening in semi-arid environments

By J. A. FRANCO*, J. J. MARTÍNEZ-SÁNCHEZ, J. A. FERNÁNDEZ and S. BAÑÓN

Departamento de Producción Vegetal, Unidad Asociada al CSIC de “Horticultura Sostenible en
Zonas Áridas” (UPCT-CEBAS), Universidad Politécnica de Cartagena, Paseo Alfonso XIII, 48,
30203 Cartagena, Spain
(e-mail: josea.franco@upct.es) (Accepted 13 August 2005)

SUMMARY
In landscaping and xerogardening projects, under semi-arid conditions, appropriate plant selection and conditioning
techniques used in the nursery during seedling production are crucial for the establishment, survival and subsequent
growth of plants after transplanting. Selecting ornamental plants with appropriate morphological and physiological
characteristics to improve nursery performance and tolerance of harsh environments is of vital importance. The use of
native species of wild flora is of increasing interest because of their capacity to adapt to adverse local environmental
conditions. However, the degree of adaptation to abiotic stresses varies considerably within a family, within a genus
and even within a species. Morphological and anatomical adaptations in seedlings include reductions in shoot height
and/or leaf area, rises in root-collar diameter and root growth potential and, often, a reduction in the shoot:root ratio.
These occur as a result of hardening and acclimation processes (pre-conditioning) during the nursery period, and are
correlated with the ability to withstand the shock of transplantation and to increase survival and plant growth
following transplantation in xerogardens and semi-arid landscapes. In addition, there are physiological characteristics
of seedlings related to osmotic adjustment and water-use efficiency, such as low stomatal conductance, leaf water
potential, leaf turgor potential and relative water content. These provide seedlings with a considerable capacity to
adapt to adverse conditions after transplantation into harsh environments. Suitable environmental conditions and
cultivation techniques in the nursery are essential to produce sturdy seedlings, with the above-mentioned
morphological and physiological characteristics. Deficit irrigation is the most commonly used pre-conditioning
technique to produce high-quality seedlings. In addition, using large-sized containers and appropriate substrates,
withholding N nutrition, inoculating arbuscular mycorrhizal fungi, applying plant growth retardants and mechanical
conditioning methods (including brushing, and shoot- and/or root-pruning) are common. Varying microclimatic
conditions (low temperature, low air humidity, enrichment with CO2, light intensity and photoperiod management) are
also used to control growth to produce high-quality seedlings with the ability to withstand transplanting shock and be
capable of rapid establishment and resumption of growth under xerogardening and semi-arid landscaping conditions.

M anaged landscapes and gardens are an intricate

blend of woody and herbaceous ornamentals,
turfgrass, organic and mineral ground cover and other
elements. Functional and sustainable landscapes and
xerogardens are created when attention is paid to
minimising or alleviating abiotic and biotic stresses from
plant selection and production to transplanting, and
finally maintenance of the established plants. Topics for
attention include the selection and use of superior taxa
for managed landscapes and xerogardens, nursery
production methods for enhanced transplant success,
establishment and post-plant maintenance protocols to
minimise stress, and intervention or rescue treatments
for established plants (Iles, 2003).
Intensive management in the nursery is primarily
geared to promote vigorous growth, whereas sub-
optimal conditions await these seedlings in the landscape
and/or xerogarden. Most natural environments, including
*Author for correspondence.
xerogardens, urban landscapes and arid and semi-arid
landscapes, are continuously sub-optimal with respect to
one or more environmental parameters such as water
availability, temperature, air humidity, soil salinity or
nutrient availability. Thus, transplantation from a nursery
into a landscape or xerogarden creates a stressful
transition period, critical to the establishment,
performance and survival of these plants. The goal of
seedling pre-conditioning is to produce sturdy plants that
have a high level of photosynthetic reserves and
adequate morphological characteristics, and therefore
are capable of rapid establishment and resumption of
growth, once in the landscape.
The environmental conditions and cultivation
techniques used in the nursery are crucial for the
establishment and subsequent growth phase in
xerogardening, re-vegetation and landscaping projects.
For example, it is known that irrigation practices affect
some morphological and anatomical aspects related to
the hardening of seedlings, and with plant growth after
4 Nursery production of ornamental plants
transplanting (Van Den Driessche, 1991a, b; Leskovar
and Stoffella, 1995; De Herralde et al., 1998; Franco et al.,
2001).
The morphological and nutritional characteristics of
plants grown under nursery conditions, and their
performance following transplantation, may be
determinants in their establishment and survival. For
example, Kailash and Kannan (1999) observed that the
higher root:shoot ratios of Cassia siamea, Peltophorum
pterocarpum and Albizia lebbeck benefited these tree
species upon planting-out in semi-arid field conditions.
Their survival rates were 92%, 84% and 90%,
respectively, whereas that of Adenanthera pavonina,
which had a low root:shoot ratio, was only 30%. C.
siamea and P. pterocarpum were also superior in biomass
production per unit of N uptake, and had a lower
nutrient demand after transplanting, so that these species
grew better under nutrient-poor field conditions.
The size, morphology and architecture of the root
system influence the relative size and growth rate of the
shoot. Optimum root systems throughout a plant’s life
cycle can ensure optimum shoot growth and
development. However, abiotic stresses that originate in
the root zone during the establishment phase can be
expressed in the shoots, thereby affecting dry-matter
partitioning and, ultimately, growth, development and
survival of the plant (Leskovar and Stoffella, 1995).
This work seeks to review the recent scientific
literature on two of the above-mentioned topics: plant
selection and nursery production methods. Substantial
research has been carried out in these areas. This
includes aspects on selection of ornamental species well-
adapted to semi-arid environments and aspects of
production methods such as container and substrate
properties, irrigation management, fertility management,
mycorrhizal inoculation, control using plant growth
regulators, mechanical conditioning, root and shoot
pruning practices and management of microclimatic
conditions. Some of the references cited are studies on
nursery production of vegetables and forestry species.
They have been included because they extend our
understanding of how techniques used during the
nursery phase may affect seedling characteristics and
performance following transplantation.
CURRENT TRENDS IN LANDSCAPING AND
GARDENING IN SEMI-ARID ENVIRONMENTS
Recent trends in landscaping and gardening under
semi-arid conditions include the use of drought-adapted
trees and shrubs in low water-use landscape designs,
while some municipalities restrict the use of turf and
offer incentives to convert to xeric designs. However,
planting schemes and maintenance practices often
include excessively dense plantings and over-irrigation of
xerophytic species, which must then be pruned
frequently to limit over-crowding and obstruction of
roads and walkways (Stabler and Martin, 2004).
The decline in traditionally designed gardens,
landscapes and urban greenspaces is a response to the
development of notions of sustainability, biodiversity and
to the de-skilling of the maintenance workforce
(Hitchmough, 2004). Alternative models usually involve
the development of ecologically-based plant communities
composed of both native and exotic species that can be
rich both ecologically and aesthetically.
Maritime climates in many regions of Europe
commonly present great opportunities and great
problems. Mild winters, when combined with a long
growing season, are favourable for a wide range of
ornamental plants, including many southern hemisphere
species which have become very popular. Arguably this
results in a planted landscape that is clichéd, repetitive
and often inappropriate (Kingsbury, 2004). Likewise, the
introduction of foreign species into desert landscapes has
caused enormous problems, altering ecological systems
and the intrinsic character of local landscapes (Kotzen,
2004).
Although native plants have largely been ignored in
greenspaces designs, the use of autochthonous species of
wild flora in xerogardening, landscaping and re-
vegetation is of increasing interest because of their
capacity to adapt to adverse environmental conditions:
hot day and low night temperatures, drought and salinity
(Zhang et al., 1996; De Herralde et al., 1998; Sánchez-
Blanco et al., 1998; Cabot and Travesa, 2000; Franco et al.,
2001; Martínez-Sánchez et al., 2003). Many of these
species are good alternatives to the ornamental species
traditionally used in semi-arid ecosystems because of
their high resistance to pests, high salt-tolerance, high
water-use efficiency and the fact that their growth patterns
are well-adapted to the prevailing environmental
conditions (Morales et al., 2000; 2001; Franco et al., 2002b;
Vignolio et al., 2002; Clary et al., 2004). It is known that
these plants have a suite of morphological and
physiological adaptations that allow them to survive
abiotic stresses; however, the degree of adaptation may
vary considerably among species, even within a species
(Sánchez-Blanco et al., 2002; Torrecillas et al., 2003).
SEEDLING TRANSPLANTATION UNDER
HARSH CONDITIONS
Amenity landscapes often require supplemental
applications of water in situations of low soil water
availability, such as in root-restricted urban settings,
during unusual or seasonal periods of low precipitation,
or in arid regions where irrigation is essential all-year-
round. Most landscape plants need short-term irrigation
following planting, until they have established new roots
into the surrounding soil. Also, as in agriculture, amenity
landscapes that have ornamental or utility value are
irrigated when rain is insufficient to support growth.
Irrigation can be permanent or temporary to
compensate for inadequate rainfall in arid and semi-arid
areas (Kjelgren et al., 2000).
Commonly, seedlings of different species irrigated
frequently following transplantation generate a greater
new root mass and establish more quickly than those
that receive infrequent irrigation, although there are
wide variations in this response (Clemens and Radford,
1986; Gilman et al., 1998; Marshall and Gilman, 1998).
Thus, root development of Limonium cossonianum
plants was substantially lower when there was no
establishment irrigation than when a single
establishment irrigation of 50 l m–2 was applied in semi-
arid conditions (Franco et al., 2002b). However,
establishment of plants in xerogardens or in arid and
 J. A. FRANCO, J. J. MARTÍNEZ-SÁNCHEZ, J. A. FERNÁNDEZ and S. BAÑÓN
semi-arid landscapes, where available irrigation water is
limited, is often costly and problematic.
On the other hand, most studies investigating the
effect of salt on plants have focussed on crop species.
However, high saline conditions adversely affect the
growth and survival of seedlings of most ornamental
species used for landscaping and gardening (Ramoliya
and Pandey, 2003). Even wild natives from littoral areas,
such as Argyranthemum coronopifolium (Morales et al.,
1998), are not salt-tolerant species. However, low salinity
has been found to stimulate germination and growth of
young seedlings in Lotus creticus (Sánchez-Blanco et al.,
1998), Pinus banksiana (Croser et al., 2001) and Protea
obtusifolia (Rodriguez-Perez et al., 2000). Nevertheless,
restoration of salt-land through re-vegetation with
halophytes can fail in the long-term because of salt
accumulation in the root-zone (Barrett-Lennard, 2002).
In California, Wu et al. (2001) evaluated the salt-
tolerance of nine landscape plants (including Pistacia
chinensis, Nerium oleander, Buxus microphylla, Nandia
domestica and Jasminum polyanthum) for recycled-
water irrigation, and presented evidence that plants with
greater salt-tolerance tended to accumulate less salt, and
that plants were more susceptible to salt-stress under
sprinkler irrigation than under drip-irrigation because of
the direct contact of leaves with salt.
In coastal gardens and landscapes, other abiotic
factors, including exposure to sea aerosols (with salt,
surfactants and other pollutants) may affect the growth
and survival of even wild native species from littoral
areas. This was the case with A. coronopifolium and
Limonium pectinatum (Sánchez-Blanco et al., 2003). The
degree of adaptation to sea aerosols may vary
considerably within a genus. For example, Cistus
monspeliensis was more tolerant to sea aerosols than C.
albidus, showing a lower reduction in plant growth and
less leaf damage, because of its ability to
compartmentalise the toxic ions intracellularly (Sánchez-
Blanco et al., 2004c).
A substantial lack of experience exists about the
suitability of ornamental plants to construct steep noise-
barrier wall systems (e.g., for motorways, airports, etc.).
Only properly-adapted species are suitable, since they
have to survive extreme temperatures, drought and,
frequently, limited root space. Furthermore, to cut costs,
low maintenance planting is required with an attractive
display at the same time (Eppel-Hotz, 2004).
When seedlings are transplanted into the field,
planting depth usually affects their survival and
subsequent shoot and root growth (Broschat, 1995).
Planting techniques also affect establishment in urban
sites, as reported by Ferrini et al. (2000).
APPROPRIATE PLANT SELECTION
There is a great need to find and use ornamental
plants that are drought- and saline-tolerant, to grow
adequately and to save water. However, the use of
foreign alien species that are heat- and drought-tolerant,
and may be salt-tolerant, is no longer universally
acceptable (Kotzen, 2004). The demand for new and
varied native ornamental plants has never been greater;
hence, programmes to develop, through selection and
hybridisation, hardy perennial plants well-adapted to
5
different climate and landscape conditions have been
initiated in the last decade (Bartual, 2000; Ault, 2003).
Also, tissue culture of native plants has been used for
mass propagation of selected native perennial plants that
cannot be propagated on a large-scale by means of seeds
and cuttings (Sudhersan et al., 2003).
Low water-use plantings may enhance water
conservation in xerogardening and dry landscapes.
However, appropriate plant selection is hindered by the
dearth of information available on the water
requirements of different plants. García-Navarro et al.
(2004) showed that measurement of water-use at the end
of nursery production for four woody species of
distinctly different ecological origin (Leucophyllum
frutescens, Spiraea vanhouteii, Viburnum tinus and
Arctostaphylos densiflora) may be useful to predict
relative water-use after their establishment in the
landscape.
Selecting ornamental plants for improved drought-
tolerance has long been of interest to those involved in
landscape horticulture, urban greenspace design and
xerogardening. Many factors contribute to drought-
tolerance, or lack of drought-tolerance in ornamental
plants, as reported by Chapman and Augé (1994) for four
native ornamental perennials widely used in gardens and
landscapes (Echinacea purpurea, Rudbeckia fulgida,
Monarda didyma and Helianthus angustifolius). In
addition, it is documented that the degree of adaptation
to drought-stress varies even within a species (Shoemake
and Arnold, 1997; Prevete et al., 2000; Reaves and
Whitlow, 2004). For example, L. creticus subsp. creticus
showed higher stomata and trichome densities on their
abaxial surface than L. creticus subsp. cytisoides. These
morphological traits caused clear differences in water-
retention of the leaf surface and in leaf reflectance. Thus,
leaf water potential and photochemical efficiency under
water-stress conditions were higher in the sub-species
creticus than in cytisoides (Savé et al., 2000). Another
example is provided by Stewart and Graves (2004). They
concluded that Rhamnus caroliniana is capable of
maintaining carbon fixation and growth over a wide
range of soil water-contents and, unlike R. cathartica, is
not dependent upon morphological, anatomical or
physiological adjustments to optimise its growth and net
photosynthesis.
It is well-documented that some wild species have a
greater degree of salt-tolerance than their cultivated
relatives. Salinity affected the ornamental aspects of
cultivated and wild Limonium spp. in different ways
(Morales et al., 2001). Thus, salinity did not affect the
ornamental quality of L. pectinatum (a wild species), in
which the production of floral stems was the same in
control and 200 mM NaCl-treated plants. However, in L.
latifolium
L. capsicum (a cultivated species), saline-
stress reduced floral stem number, height and dry weight
(DW) compared to control plants. Similarly, C. albidus
and C. monspeliensis plants, irrigated with water of
different salinity, developed avoidance and tolerance
mechanisms at morphological and physiological levels
(i.e., reduction in canopy area, Na+ and Cl– inclusion and
osmotic adjustment). Nevertheless, the reaction of each
species to osmotic adjustment was different. C.
monspeliensis plants showed a greater capacity to absorb
water and were able to conserve it more efficiently than
6 Nursery production of ornamental plants
C. albidus plants (Torrecillas et al., 2003). Also, in follow-
up work, Sánchez-Blanco et al. (2004c) determined that
C. albidus was more sensitive to sea aerosols than C.
monspeliensis, showing greater leaf damage and
markedly decreased growth.
The daily light integral usually affects growth and
flowering of ornamental plants, as reported by Fausey
et al. (2005) for Achillea millefolium, Gaura lindheimeri
and Lavandula angustifolia. Tolerance to high light-
intensities during nursery production, and subsequent
establishment in the landscape, may vary within a genus
and within a species (Heiskanen, 2004). However, in five
Illicium species, different parameters (viz. CO2
assimilation rate, dark respiration rate, chlorophyll and
total carotenoids contents) improved under low light,
suggesting that optimal growth occurs for all taxa in
shaded conditions (Olsen et al., 2002). An extended
photoperiod and light-quality affected growth of
Carpinus caroliniana, Fagus grandifolia and
Gymnocladus dioicus seedlings (Richardson-Calfee
et al., 2001). Similarly, in a recent review on photoperiod
and plant growth, Adams and Langton (2005) have
shown that long-day treatments usually promote an
increase in DW in a number of ornamental plants that
otherwise grow in short-days.
Over-Winter survival ratios of various ornamental
grasses have been reported by Meyer and Cunliffe
(2004), who found significant differences between
species within the Miscanthus genus. Schrader and
Graves (2004) also found differences in cold-acclimation
for the three Alnus maritima sub-species. Similarly, in a
recent review article by Wilson and Struve (2004), a
perspective on over-Winter mortality in stem cuttings
was developed, reporting different degree of
susceptibility to Winter injury between genera, but also
within a genus. Therefore, substantial differences in cold-
hardiness should be considered when selecting plants for
landscaping purposes outside their native range.
During vegetative reproduction, the influence of
cutting position on the mother stem on rooting ability
has been widely documented, although the exact effect
differs from species-to-species. In Protea spp.
(Montarone et al., 1997) and Dorycnium spp. (Alegre
et al., 1998), the best results were obtained with apical
cuttings, while in Rosa centifolia both apical and sub-
apical cuttings produced equally good results (Al-Saqri
and Alderson, 1996). In N. oleander, basal cuttings
produced larger root growth, although more roots with a
more homogeneous length distribution were obtained
from apical cuttings (Ochoa et al., 2003).
To conclude, to improve landscape restoration and
xerogardening, it is necessary to know the mechanisms of
plant responses to environmental factors. It is also
necessary to understand acclimation processes during
nursery production or during the first stages after
transplantation. The degree of adaptation to abiotic
stresses may vary considerably within a family, within a
genus and even within a species. Therefore, appropriate
selection may improve both the nursery and landscape
performance of plants (Savé et al., 2000; Torrecillas et al.,
2003; Sánchez-Blanco et al., 2004c; Sharma and Graves,
2004; Schrader and Graves, 2004; Shoemake et al., 2004).
Web-sites may serve a useful role in the selection of
ornamental plants (Lehrer and Brand, 2003).
NURSERY PRODUCTION AND CONDITIONING
Container size and type
The issue of container size is extremely important
both to seedling producers (i.e., nurseries) and seedling
users. A trend in many nurseries is toward smaller
containers, thereby reducing the need to use more space.
This trend also reduces production cost per plant. While
the use of smaller containers may improve the efficiency
of seedling production, it is unclear how plants grown
under conditions of increased root-restriction will
perform after transplanting into diverse environmental
conditions.
Container size can affect the growth and development
of bedding plants. Thus Van Iersel (1997) observed that
Salvia splendes plants grew faster in 510 ml pots than
those in 55 ml or 166 ml pots, because of their higher leaf
area ratio. Both lateral branching and leaf expansion
were suppressed by root restriction. Container size also
influenced the growth of five ornamental grasses
(Schizachyrium scoparium, Sporobulus heterolepis,
Calamagrostis acutiflora and two Miscanthus spp.)
during nursery production, but had no effect on over-
Wintering survival (Meyer and Cunliffe, 2004). In
addition, seedling establishment and survival in semi-
arid environments were improved when larger
container sizes were used in conjunction with
appropriately prepared sites (NeSmith and Duval, 1998;
Pastor et al., 1999; Loveall et al., 2002). In contrast, using
different container sizes, no differences were observed
in root and shoot growth in the nursery, or in survival
rates after transplantation of Shorea leprosula (Aminah
et al., 2004).
On the other hand, container-type may affect root
system characteristics during nursery production. A
review by Appleton (1993), with special reference to
ornamental woody plants, studied several nursery
production alternatives to reduce or eliminate “circling”
roots. Woody plant roots that “circle” during nursery
production have the potential, once transplanted, to
enlarge and becoming “girdling” roots that may stress or
kill the plant. Alternatives studied have included the use
of different container types, such as in-ground fabric,
rigid plastic, ‘pot-in-pot’, above-ground modified rigid
plastic, low-profile or copper-treated containers. Also,
different container-types, including standard black
plastic, low-profile plastic, fabric, wooden boxes,
chemical root-pruning or air root-pruning containers,
have different abilities to reduce the number of roots
deflected by the container wall (Marshall and Gilman,
1998). These also show different levels of effectiveness in
stopping root extension growth at the container wall-
growing medium interface (Marler and Willis, 1996) and
affect shoot and root growth (Harris and Gilman, 1993).
Substrate properties
Commercial substrates used for seedling production
and for rooting cuttings in nurseries have different
physical and chemical properties which may modify
rhizogenesis, root growth and the characteristics of the
root system. Studies on the effect of substrates on
rhizogenesis and on the morphological, anatomical and
physiological characteristics of root systems attribute
great importance to aeration, oxygen diffusion and easily
available water content (Gopikumar and Minichandran,
 J. A. FRANCO, J. J. MARTÍNEZ-SÁNCHEZ, J. A. FERNÁNDEZ and S. BAÑÓN
2002; Ochoa et al., 2003). Usually, lightweight potting
media such as coconut husk and fibre dust, peat, wood
fibre, bark compost, decomposed oil palm mesocarp
fibre or rice hulls ensure aeration, increase root and
shoot growth and can reduce the total amount of
irrigation water used, as well as greatly facilitating the
transportation of seedlings to planting sites (Aminah et
al., 2004; Bohne, 2004).
Hinesley et al. (1999) observed that peat-amendment
substrates improved seedling growth in the nursery and
subsequent field performance. They reported that small,
but significant, residual effects of peat amendments were
still evident after one growing season in the field.
Similarly, Pastor et al. (1999) concluded that, after
transplantation, better development was obtained in V.
tinus and Spiraea japonica seedlings grown in containers
filled with coarse rather than fine pine bark. The latter
plants were better adapted to cope with water-stress.
Conversely, in establishment trials of ten species of
ornamental trees and shrubs, Clemens and Radford
(1986) observed that survival was unaffected by the
potting medium. More recently, other authors (Aminah
et al., 2004; Meyer and Cunliffe, 2004) have also
determined that the media used in nursery production
do not significantly affect survival rates after
transplantation.
Linderman and Davis (2003) found that different
potting mixes (peat/coconut fibre dust) affected growth
in several ornamental plants (Tagetes patula, Teucrium
fruticans and L. angustifolia) inoculated with arbuscular
mycorrhizal fungi. Plant growth was depressed in media
highly amended with coconut-dust, with or without
arbuscular mycorrhizal fungal inoculation, although
there were no detrimental effects of the coconut-dust on
mycorrhiza formation.
Root plugs may serve as immediate sources of
nutrients for the growth of newly planted seedlings,
enhancing their early field performance, and benefiting
initial establishment of the plantation (Idris et al., 2004).
Furthermore, with this method it is possible to produce
seedlings with large fibrous root systems and a field
performance similar to other methods, but more
economically (McCreary and Lippitt, 2000).
Irrigation management
The differential provision of water to seedlings
influences their morphology, physiology and dry matter
partitioning between roots and shoots. Hence, irrigation
management relative to modification of seedling growth
is of the utmost importance for the nursery manager in
order to produce high-quality seedlings with the ability to
survive and grow better following transplantation under
different environmental conditions (Leskovar, 1998).
The application of a pre-conditioning drought
treatment during the last months of nursery culture, or
throughout nursery production, is one potential
technique to reduce the shock of transplantation that is a
key hurdle to plantation success in the Mediterranean
climate. However, drought-stress, the most commonly
used hardening treatment, can decrease photosynthetic
rates, delay flowering and increase levels of endogenous
inhibitors if the treatment is prolonged or too severe.
Crop coefficients (Kc) for containerised woody
ornamental plants are considerably higher than those
7
reported for field crops (Schuch and Burger, 1997).
Knowledge of Kc values is of great interest as nurseries
attempt to conserve water, reduce run-off and pollution
of groundwater, and establish irrigation-deficit
treatments to increase the subsequent drought-
resistance of seedlings. The severity of any water
restriction is critical. A desirable level of deficit-
irrigation results in stocky, stress-resistant seedlings; but,
if the water restriction is too severe, seedlings die or are
over-hardened, slowing new shoot and root growth
(Liptay et al., 1998).
Therefore, monitoring nursery moisture regimes and
understanding morphological and physiological shoot
and root responses of seedlings to water management
throughout the nursery period are critical to optimise
production of high-quality seedlings.
Numerous reports have determined that deficit-
irrigation during nursery production affects some
morphological and physiological aspects related to the
hardening of seedlings, or plantlets, of ornamental plants of
interest for xerogardening, re-vegetation and landscaping,
plants such as A. coronopifolium (De Herralde et al., 1998),
L. cossonianum (Franco et al., 2002b), L. creticus (Franco
et al., 2001; Bañón et al., 2004), Myrtus communis (Bañón
et al., 2002), N. oleander (Bañón et al., 2005b), Olea
europaea var. sylvestris (Bañón et al., 2003b), Phillyrea
angustifolia (Fernández et al., 2004; 2005), Pistacia lentiscus
and Juniperus oxycedrus (Vilagrosa et al., 2003), Rhamnus
alaternus (Bañón et al., 2003c) and Rosmarinus officinalis
(Sánchez-Blanco et al., 2004b).
Deficit-irrigation during the nursery phase reduced the
growth parameters of the aerial parts (i.e., plant height,
shoot length, leaf area, stem diameter, shoot fresh and
DW) and root growth (i.e., root length, DW and volume)
of Acer pseudoplatanus (Hipps et al., 1996), Cotinus
coggygria, Forsythia intermedia (Cameron et al., 2004), L.
creticus (Franco et al., 2001; Bañón et al., 2004), M.
communis (Bañón et al., 2002), N. oleander (Bañón et al.,
2005b) and R. officinalis plants (Sánchez-Blanco et al.,
2004b). With deficit-irrigation, the shoot:root ratio fell by
0.20 for R. officinalis, by 0.33 for N. oleander, by 0.36 for
A. pseudoplatanus, by 0.45 for L. creticus and by 0.71 for
M. communis.
In addition, deficit-irrigation during the nursery phase
increased the percentage of thick roots, and reduced the
percentage of medium and fine roots in M. communis
and N. oleander plants (Bañón et al., 2002; 2005b).
Proportionally, root volume was reduced more than DW,
with the result that root density increased under deficit-
irrigation. The hardening of roots, as revealed by an
increased percentage of brown roots at lower irrigation
rates in L. creticus (Franco et al., 2001) and L.
cossonianum (Franco et al., 2002b), is of great interest to
produce seedlings that are better adapted to drought-
stress at transplantation. The change in colour from
white to brown is associated with suberisation of the
exodermis, and may reflect a metacutisation process. This
is a process of lignification and suberisation that results
in a resting root being protected against significant
fluctuations in environmental conditions such as
drought, and being capable of re-growth when conditions
ameliorate (Bloomfield et al., 1996).
At the end of the nursery period, the stomatal
conductance (gs), leaf water potential (l), leaf turgor
8 Nursery production of ornamental plants
potential (p), leaf osmotic potential at full turgor (os),
and relative water content (RWC) of N. oleander, P.
lentiscus, Quercus coccifera, J. oxycedrus, R. officinalis
and M. communis plants grown with deficit-irrigation
were lower than those of normally-irrigated plants
(Bañón et al., 2002; 2005b; Vilagrosa et al., 2003;
Sánchez-Blanco et al., 2004b). Deficit-irrigation
produced L. creticus seedlings with greater leaf
trichome densities and numbers of xylem vessels in
stems and roots, and induced a range of physiological
adaptations. In addition, seedlings undergo osmotic
adjustment that might provide considerable capacity to
adapt to adverse conditions during the field
establishment period (Franco et al., 2002a; Bañón et al.,
2004). The above-mentioned changes allow plants to
maintain higher net CO2 assimilation rates and an
elevated photosystem II status, thus facilitating an
increase in drought survival (Vilagrosa et al., 2003).
Also, P. angustifolia plants watered at 40% water-
holding capacity throughout the nursery phase showed
higher stomatal density and water-use efficiency, which
would allow the plants to overcome transplantation
shock (Fernández et al., 2004).
C. albidus and C. monspeliensis plants, submitted to
irrigation water-deficit, developed avoidance
mechanisms based on stomatal closure, epinasty and
reductions in leaf area and root hydraulic conductivity.
After the water-deficit, these features contributed to
increase relative growth and net CO2 assimilation rates
during the recovery period (Sánchez-Blanco et al.,
2002). Differences in drought responses of Boltonia
asteroides, Eupatorium rugosum and Rudbeckia triloba
(ornamental herbaceous perennials, and members of
the Asteraceae) were attributed to differences in water-
use rates. In turn, those may be attributed to many
factors, such as differential growth of roots, shoots and
leaves, changes in carbohydrate partitioning and gs
(Prevete et al., 2000).
Deficit-irrigation of Prosopis glandulosa (Bainbridge
et al., 2001) and N. oleander (Bañón et al., 2005b)
seedlings during nursery production clearly reduced
their mortality rates after transplantation under drought
and heat conditions. Similarly, Hipps et al. (1996) found
that, after transplantation, shoot growth in A.
pseudoplatanus seedlings that had previously received a
higher rate of irrigation was less than in those that
received a lower rate. Equally, seedlings of L. creticus
(Franco et al., 2001; 2002a) and L. cossonianum (Franco
et al., 2002b) that were watered less in the nursery
showed greater and faster root development following
transplantation under semi-arid conditions, especially
when soil moisture was low (Franco et al., 1999). This
adaptive potential of plants perseveres, and a small
amount of water after a long period of drought
reactivated root growth more rapidly in L. cossonianum
plants that had been less watered during the nursery
period (Franco et al., 2002b). Furthermore, low irrigated
L. creticus plants in the nursery had greater post-planting
stem growth over the same soil surface area covered, and
therefore provided more dense cover. Similarly, the
aerial parts of L. cossonianum plants hardened during
the nursery phase developed better, had a more compact
appearance, and had longer flowering stems. In L.
creticus, the harsher the conditions after transplanting
(i.e., less water during establishment irrigation), the more
evident was the positive effect of hardening in the
nursery (Franco et al., 2001).
Nevertheless, Picea mariana seedlings, subjected to
four irrigation regimes [from 15% to 60% (v/v),
water/substrate], were unaffected by substrate water
content, showing no differences in height, stem diameter,
number of needle primordia or root hydraulic
conductivity. Consequently, it is possible that reducing
the quantity of irrigation water used during the nursery
period does not significantly affect the growth and other
characteristics of seedlings (Bergeron et al., 2004).
On the other hand, the irrigation system can influence
root growth, resulting in differences in lateral and basal
root elongation (Leskovar and Stoffella, 1995; Franco
and Leskovar, 2002), shoot water potential (Leskovar,
1998) and mineral nutrient content (Biernbaum and
Versluys, 1998; Franco and Leskovar, 2002). Traditionally,
ornamental seedlings for gardening are grown with
overhead irrigation. However, sub-irrigation (flotation
or ebb and flow) and other systems, including capillary
wick containers, capillary mats and water collection
reservoirs beneath individual containers, can produce
high-quality seedlings for xerogardening and
landscaping. They also minimise negative environmental
impacts by reducing the water use and the load of
nutrients applied to the ground (Bilderback, 2002;
Yeager and Henley, 2004). However, these systems do
not significantly reduce losses of nutrients or pesticides,
unless used in conjunction with a recirculation system
(Harris et al., 1997).
Fertiliser management
One of the most effective ways to control seedling
growth during nursery production is to moderate the
nutritional regime. Consequently, a number of studies
have been conducted to evaluate the relationship
between fertility practices during nursery production
and plant establishment, growth and performance after
transplantation into xerogardens or landscape sites with
minimum management. Adequate growth of seedlings
under different climates is promoted through the
application of appropriate fertilisation, resulting in a
balanced growth in terms of shoot:root ratio, which is
vital for plants growing in harsh environments where
competition for water and nutrients is usually intense
(Martin and Ruter, 1996; Mexala et al., 2002; Rikala et al.,
2004).
Cabrera and Devereaux (1999) concluded that, 4
months after transplantation into the landscape, plant
biomass was higher in Lagerstroemia indica
L. fauriei
plants that had been grown with higher N supplies
during the nursery period, and had been among the
smallest seedlings at transplantation. Plant shoot:root
ratio and tissue N, Ca, S and Fe concentrations, which
had been affected by the N fertilisation regime in the
nursery, equalised after transplantation, with no
significant differences observed between treatments 4
months after transplantation. Likewise, plant survival
was not affected by treatment. However, the flowering
response in the landscape was delayed in plants grown in
the nursery with higher N supplies. Similarly, Ilex cornuta
seedlings supplied with higher N during nursery
production gained more new shoot weight, but less root
 J. A. FRANCO, J. J. MARTÍNEZ-SÁNCHEZ, J. A. FERNÁNDEZ and S. BAÑÓN
weight, after transplanting into a landscape situation,
than seedlings that had received less N (Gilman et al.,
1996). Another example is provided by Rikala et al.
(2004), who reported that, by improving the pre-planting
nutrient status of seedlings during nursery production,
shoot growth, stem diameter and root biomass can be
enhanced after transplantation, especially into nutrient-
poor soils.
The effects of fertilisation can vary within a genus.
Close and Beadle (2004) observed that Eucalyptus
globulus had greater foliar concentrations of N and P at
planting than E. nitens after nursery production under
the same fertiliser regime. They concluded that there was
nutrient retranslocation associated with growth after
transplantation, and observed a decrease in N, and
generally P concentrations in the leaves. Despite
differences in N and P concentrations, partitioning of
total N to soluble nucleic acids and proteins, and of total
P to nucleic acids, sugars and insoluble P were similar
between high- and low-nutrient treatments during
nursery production.
As explained below, mycorrhizal fungus-inoculated
seedlings may improve establishment by enhancing the
growth and nutrition of newly-planted seedlings.
However, successful nursery colonisation of planting
stock by these fungi is often incompatible with
conventional fertiliser application practices because of
their toxic or inhibitory effects (Quoreshi, 2003).
It can therefore be concluded that fertiliser
application usually has only a minor effect on seedling
morphology, but that high N supplies produce high
shoot:root ratios in a number of species, thus prejudicing
their subsequent growth and survival after planting-out
in semi-arid environments.
Mycorrhizal inoculation
The early development of a functional root system is
essential for the successful establishment of many
ornamental woody species, particularly in adverse
environmental conditions, where large variations in the
ability to obtain water and to acquire nutrients by
arbuscular mycorrhizal fungi (AMF)-inoculated root
systems have been reported (Goicoechea et al., 2004;
Iglesias et al., 2004; Sánchez-Blanco et al., 2004a). It has
been observed that AMF-infection can induce
morphological changes to the root system of the host
plant such as differential root branching. Morphometric
analyses of roots of N. oleander, O. europaea, O.
europaea var. sylvestris, P. lentiscus, Populus spp., Retama
sphaerocarpa, Rhamnus lycioides and R. officinalis
showed that AMF-plants developed a denser root
system, with larger numbers of short, branched roots,
which positively affected the establishment of seedlings
and plantlets (Hooker et al., 1992; Citernesi et al., 1998;
Palenzuela et al., 2002; Marín et al., 2003; Sánchez-Blanco
et al., 2004a).
The effect of AMF on drought-resistance of host
plants has been studied in several wild plant species of
increasing interest in xerogardening and landscaping.
The response of mycorrhizal plants to drought-stress
depends on the fungal species, on interactions between
the plant species and the introduced fungus (Sánchez-
Blanco et al., 2004a) and on the degree of drought-stress
(Savé et al., 1994).
9
Anthyllis cytisoides, a shrub legume used to recover
desertified semi-arid areas, is highly colonised by AMF in
natural conditions, as mycorrhizal symbiosis is crucial for
their growth, and P and N nutrition. In a recent study
with container-grown Ipomoea carnea subsp. fistulosa,
Carpio et al. (2005) also found greater nutrient utilisation
in AMF-inoculated plants than in non-inoculated plants.
Goicoechea et al. (2004) suggested that is possible to
improve the functionality of the symbiosis of A.
cytisoides with Glomus fasciculatum and Rhizobium by
applying low N and P fertilisation during the nursery
period. Furthermore, according to the functionality of
both micro-symbionts, the most appropriate moment to
transplant A. cytisoides from the nursery to natural
conditions would be between the third and fourth month
after sowing. Iglesias et al. (2004) found that, 9 months
after transplantation, P, K, Ca, Mg, Cu and Zn uptake
levels were greater in AMF-inoculated Taxus baccata
plants than in those not inoculated during nursery
production. However, this has not always been the case.
AMF-inoculation has shown no apparent benefit during
nursery production and initial establishment in other
experiments (Gilman, 2001; Linderman and Davis, 2003;
Martin et al., 2003), usually in years with below-average
rainfall and a severe Summer drought (Maestre et al.,
2002).
Carpio et al. (2003) studied the effects of commercial
AMF on the growth, survival and subsequent landscape
performance of several ornamental plants (Acacia
greggii, Chilopsis linearis, Diospyros virginiana, Platanus
occidentalis, I. carnea and Plumbago auriculata) grown in
a nursery. They found that AMF enhanced seedling
growth during nursery production. At the end of the first
growing season after transplantation, AMF-inoculated P.
occidentalis and C. linearis plants showed more growth
than non-inoculated plants. Also, AMF-inoculated P.
auriculata had higher survival rates than non-AMF-
inoculated plants. However, by the end of the second
growing season, there were no differences in growth or
survival between AMF-inoculated and non-inoculated
plants.
AMF-colonisation in the nursery is maintained after
transfer to the final habitat, as reported in Tilia cordata
by Nielsen and Rasmussen (1999) and in T. baccata by
Iglesias et al. (2004).
In the above research, AMF-inoculation was carried
out with Glomus intraradices (Palenzuela et al., 2002;
Carpio et al., 2003; 2005; Linderman and Davis, 2003;
Marín et al., 2003), G. mosseae (Citernesi et al., 1998;
Marín et al., 2003), G. deserticola (Iglesias et al., 2004;
Sánchez-Blanco et al., 2004a), G. fasciculatum
(Goicoechea et al., 2004), a commercial AMF inoculum
containing a mix of Glomus spp. (Carpio et al., 2003;
2005), Acaulospora scrobiculata (Iglesias et al., 2004) or
Pisolithus tinctorius (Maestre et al., 2002; Martin et al.,
2003).
In conclusion, AMF-inoculation can enhance growth
during nursery production. AMF-inoculated plants have
improved water relations, greater tolerance to
environmental stress and better survival after
transplanting when compared to non-inoculated plants.
But this is not always the case, mycorrhizal inoculation
does not always show benefits during the nursery phase
or subsequent transplantation.
10 Nursery production of ornamental plants
Control using plant growth regulators
Plant growth regulators (PGRs) have been used to
manipulate the shape, size, flowering and abiotic stress
tolerance of ornamental plants (Ruter, 1994; Bañón et al.,
2001b). PGRs can also influence the differential
development behaviour of roots, buds and shoots during
clonal propagation by in vitro techniques, as shown by
Glocke et al. (2005) in ornamental eucalypts.
Paclobutrazol (PBZ), a growth retardant and member
of the triazole group, induces mild water-stress tolerance
and can increase water-use efficiency in seedlings and
adult plants (Van Den Driessche, 1996; Watson, 2001).
PBZ-induced drought-tolerance has been associated
with decreases in transpiration, plant height, biomass and
leaf area, and with an increase in stomatal resistance in
nursery-produced P. angustifolia seedlings (Fernández et
al., 2004; 2005). These effects increased the percentage of
plants that survive after transplantation into semi-arid
conditions (Bañón et al., 2001b). Similarly, PBZ reduced
plant height and aerial DW, increased root diameter and
root system volume, and reduced stomatal conductance
in Arbutus unedo (Navarro et al., 2004).
PBZ reduced the symptoms of saline-stress in N.
oleander seedlings by reducing the uptake and
accumulation of harmful Na+ and Cl– ions by plant
tissues, and promoted a process of osmotic adjustment
through the accumulation of organic osmotic compounds
(Bañón et al., 2005a). Also, PBZ reduced the symptoms
of saline-stress and mortality in R. alaternus seedlings in
different ways; by increasing stomatal conductance, by
promoting organic solute synthesis and by reducing the
availability of salt ions in the medium (Bañón et al.,
2003d).
Application of PBZ to Pyracantha sp. and Juniperus
chinensis during nursery production reduced the total
biomass of seedlings, which can be correlated with an
ability to withstand the shock of transplantation. After 9
months in the landscape, PBZ continued to influence
plant height, plant width and shoot DW in Pyracantha
sp., but had no effect on J. chinensis (Ruter, 1994).
Chlormequat chloride (CCC) reduced the DW of the
aerial parts of N. oleander plants grown in a nursery,
thereby producing plants with a darker and duller colour
(Bañón et al., 2001a). Similarly, both PBZ and ethephon
(ETH) controlled stem height in Reichardia tingitana
and improved its ornamental value (Bañón et al., 2003a).
A mix of daminozide and CCC effectively reduced the
height of six perennial bedding plants including
Gaillardia grandiflora, Salvia greggii and Heliotropium
arborescens (Latimer et al., 1999). However, these
responses did not persist after transplantation into the
landscape. Only G. lindheimeri, previously treated with
uniconazole, exhibited persistent growth reduction 8
weeks after transplantation (Latimer et al., 1999).
Different PGRs can have very different effects, as shown
in two studies by Bruner et al. (2000; 2001) who
determined the effects of daminozide, flurprimidol,
uniconazole, PBZ, CCC and ETH on vegetative growth
and flowering in Canna
generalis during nursery
production and landscape establishment.
In short, the above studies show that PGRs, mainly
PBZ, can be used widely during nursery production to
improve seedling tolerance to abiotic stresses, and to
increase plant survival rates during establishment in
harsh conditions. PGR-induced stress-tolerance does not
always persist after transplantation.
Mechanical conditioning
Mechanical stresses, including shoot brushing, shaking
of potted plants, vibrating pots or plants, wind or forced-
air treatment, and stem rubbing or flexing, have been
proposed as alternative conditioning treatments for
seedling production, because they improve physical stem
strength and stress-tolerance. Nevertheless, each species
responds differently to these treatments (Garner et al.,
1997; Latimer, 1998). Brushing seedling shoots has
probably received most attention from researchers
working with vegetable seedlings, or ornamental
perennial and annual plants. Brushing provides tactile or
thigmic stimulation of the growing points of the seedling.
The heights of several ornamentals, including
Callistephus chinensis, Dendranthema grandiflorum,
Senecio cineraria, Tagetes erecta and Petunia
hybrida,
have effectively been reduced by brushing; but Ageratum
houstonianum was not responsive to this technique.
Plant growth habit appears to affect seedling response to
mechanical condition by brushing. More brittle or stiff
seedlings undergo less bending action than seedlings
with more flexible stems, and thereby exhibit less
reduction in plant growth (Latimer and Oetting, 1997;
Latimer, 1998).
Pruning
Frequently during nursery production, air or
chemically root-pruned seedlings survive and grow
better following transplantation (Gilman, 2001; Pardos
et al., 2001; Krasowski, 2003). Conifer seedlings grown in
copper-treated polybags had greater height, root collar
diameter, biomass production, higher root:shoot ratios
(because of less root egression) and less root-spiralling
than non root-pruned seedlings. Therefore, this
production system improves seedling quality and
consequently improves survival and field performance
after transplantation (Aldrete and Mexal, 2002).
In Betula pendula, progressive root removal prior to
planting promoted changes in water relations, and
reduced shoot and root growth under well-watered
conditions. Root re-growth was proportional to the
amount of old root material left behind after
transplantation (Bellett-Travers et al., 2004). Similarly,
Andersen et al. (2000) determined that pruning the root
system before transplanting reduced re-growth in
accordance with the severity of pruning, with shoots
being more affected than roots. Removal of coarse roots
depressed final root DW, whereas removal of fine roots
reduced shoot DW and hence shoot:root ratio. Hipps
et al. (1996) also found that A. pseudoplatanus seedlings,
that had previously been root-pruning in the nursery,
had greater lengths of fine roots and root:shoot ratios
than those not root-pruned. However, there was no
significant effect of root-pruning on shoot growth after
planting-out.
With regard to the aerial parts, “pinching” N. oleander
plants in the nursery produced shorter plants with more
shoots, greater root length and lower shoot:root ratios
(Bañón et al., 2001a). Likewise, frequent pruning after
transplanting lowered the water-use efficiency of well-
irrigated N. oleander and L. frutescens shrubs, whereas
 J. A. FRANCO, J. J. MARTÍNEZ-SÁNCHEZ, J. A. FERNÁNDEZ and S. BAÑÓN
water-use efficiency was higher for unpruned shrubs
receiving low irrigation (Stabler and Martin, 2004).
Temperature management
Modification of nursery temperature regimes can
influence the performance of nursery stock after
transplantation into the field. The direct effects of
temperature on seedling growth are well-known, but the
after-effects of temperature on plant growth in
subsequent months, in the new planting environment,
have not been studied extensively. At the end of the
nursery phase, a decrease in nursery temperature
reduced plant height, height:diameter ratio and
shoot:root ratio in Picea glauca, Pinus contorta and
Pseudotsuga menziesii (Van Den Driessche, 1991a).
Similarly, lowered temperatures reduced shoot length
and the shoot:root ratio, and increased the percentage of
brown roots in L. creticus (Franco et al., 2001).
Nursery temperature can affect gs and transpiration
(E) after planting. Low temperatures reduced gs and E
(adjusted for xylem pressure potential) and specific leaf
area in P. glauca, P. contorta and P. menziesii during the
nursery phase, but no effect on gs and E could be
detected 12 weeks after transplantation (Van Den
Driessche, 1991b). Similarly, low temperatures increased
l and the abaxial stomatal density of water-stressed L.
creticus plants (Bañón et al., 2004). Photochemical
efficiency decreased with temperature during nursery
production of E. globulus and E. nitens, increasing foliar
anthocyanin concentrations (Close et al., 2004). Seedlings
high in anthocyanins were associated with low gs and low
specific leaf area before planting. It was concluded that
the concentration of foliar anthocyanins in the nursery is
a potential indicator of seedling hardiness to low
temperatures experienced after transplantation of
eucalypts.
Low nursery temperatures continue to affect growth
after planting, increasing relative growth rates. Survival
of transplanted P. glauca, P. contorta and P. menziesii
seedlings was inversely related to nursery temperature
(Van Den Driessche, 1991a). A similar effect was also
found in L. creticus, with plants from an unheated
greenhouse showing slightly higher root growth than
plants from a heated greenhouse after transplanting with
different levels of establishment irrigation (Franco et al.,
2001).
When the combined effects of deficit-irrigation and
low temperature during the nursery period are studied
(Franco et al., 2001; 2002a; Bañón et al., 2004), regimes
involving the least water and lowest temperatures
produce plants best adapted to stress during
transplantation. These plants have lower shoot:root
ratios, lower shoot fresh weight:length ratios, higher
percentages of brown roots, lower gs, l, p, and RWC
values, higher leaf trichome densities and greater
numbers of xylem vessels in stems and roots. The most
stressed seedlings in the nursery (i.e., those that received
least water and low temperatures) show greater and
more rapid root growth after transplantation, especially
when soil moisture content is low.
Management of air humidity
The influence of ambient air humidity in the nursery
on pre-conditioning of seedlings has received limited
11
attention. Bañón et al. (2003c), working with R. alaternus,
reported that low air humidity and high water-deficit
during the nursery phase reduced shoot growth, l, leaf
gs and net CO2 assimilation rate, leading to improving the
survival of seedlings at the end of the establishment
period. The effect of low air humidity on l of O.
europaea var. sylvestris was less marked than the effect of
deficit-irrigation. However, p values were higher,
suggesting that low air humidity pre-conditioning might
trigger an adjustment mechanism which enhances cell
wall elasticity and avoids any substantial reduction in the
leaf turgor in this species (Bañón et al., 2003b). The
combined effect of both hardening treatments increased
the rate of survival after transplantation by around 63%
compared with the control (0%).
In a subsequent study, the combined effects of deficit-
irrigation and low air humidity during the nursery phase
also reduced the mortality rate of N. oleander seedlings
after transplantation under drought and heat conditions
compared with controls (from 93% to 32%; Bañón et al.,
2005b). Similarly, the above-mentioned combined effect
reduced the mortality rate of M. communis seedlings
after transplantation under drought and heat conditions
from 100% to 66%, compared with the controls (Bañón
et al., 2002). Such behaviour was related to
morphological changes observed in the aerial parts (i.e.,
smaller plant size and lower leaf area), in the roots (i.e.,
shorter, thicker, more dense and less ramified) and in the
shoot:root ratio (i.e., reduced by approx. 60% in both
species) of the pre-conditioned plants. This behaviour
was also related to physiological changes such us the
development of an osmotic adjustment, efficient
stomatal regulation and reordering of the assimilate
gradient as the flow of solutes towards the roots
intensified. Likewise, Sánchez-Blanco et al. (2004b)
found that R. officinalis plants that had been exposed to
deficit-irrigation and low humidity pre-conditioning in
the nursery showed more efficient stomatal regulation
(lower gs values) and developed a leaf osmotic
adjustment, which were maintained during the
transplanting period. After transplantation, and during
the establishment period, these plants had a better water
status (higher l and gs values) and a greater survival
rate (43.8%) compared with the controls (0%).
As indicated in these studies, the combined
application of low air humidity and deficit-irrigation
produced a range of hardening in seedlings during the
nursery phase. This apparently resulted from a
combination of three important mechanisms for
acclimation to drought and heat: i) reduction in growth,
accompanied by a shift in the allocation of assimilates
from shoots to roots; this would reduce transpiration and
favour increased development of an efficient root
system; ii) leaf osmotic adjustment, permitting the
maintenance of leaf turgor even when soil water
availability was low through the absence of irrigation;
and iii) more efficient stomatal regulation, avoiding
excessive water loss through transpiration when soil
water levels are limiting.
Air enrichment with CO2
The use of enriched CO2 atmospheres during nursery
production is a promising technique to increase
productivity and to obtain seedlings with higher
12 Nursery production of ornamental plants

root:shoot ratios, which correlate with the ability to
withstand transplanting shock under water-stress in
Mediterranean-type ecosystems, as reported by several
studies (Wullschleger et al., 2002; Biel et al., 2004; Cortes
et al., 2004). Seedlings of Quercus cerroides and Q. ilex
increased their total biomass by approx. 60% in response
to CO2-enrichment between 500–700 µl l–1. This increase
was due entirely to an elevated root biomass, as reported
by Cortes et al. (2004). Biel et al. (2004) following up this
study, working with Pinus nigra seedlings, and analysed
the combined effects of water-deficit and air CO2-
enrichment in the nursery as a route to increase
hardening and vigour of seedlings and their performance
after transplantation. CO2-enrichment increased total
biomass, leaf biomass and the leaf area of seedlings,
while water-deficit decreased leaf area, leaf biomass and
stem biomass, without any significant interaction
between the two factors. After transplantation, previous
CO2 and water-deficit treatments did not affect plant
survival.
DISCUSSION
As indicated in this review, appropriate plant
selection, microclimatic conditions and cultivation
techniques used in the nursery during seedling
production are crucial for plant establishment, survival
and subsequent growth after transplantation. However,
in landscaping, re-vegetation and xerogardening
projects, high plant mortality rates can occasionally occur
post-planting due to poor seedling viability at the time of
transplantation, mainly under harsh environmental
conditions.
Consequently, measurements of the morphology, size
and growth potential of seedlings at the end of the
nursery period are useful because of their ability to
identify low-vigour or damaged plants that will perform
poorly when transplanted into the landscape or
xerogarden. Parameters that can be measured include:
stem height, stem flexibility, shoot growth potential, leaf
area, root-collar diameter, total plant DW, shoot:root
ratio, stem height:root-collar diameter ratio, root weight
ratio and root growth potential (Percival, 2004). Plant
survival frequently shows a significant negative
correlation with stem height, leaf area, stem
height:diameter ratio and shoot:root ratio (Table I). A
reduction in the shoot:root ratio (caused either by
reducing the shoot growth rate, or by increasing root
growth relative to shoot growth) as a result of hardening
and acclimation processes during the nursery period (i.e.,
pre-conditioning), increases plant growth and reduces
the mortality rate after transplantation under drought
and heat conditions. In addition, some physiological
characteristics of seedlings related to osmotic adjustment
and water-use efficiency (gs, l, p and RWC) might
provide considerable capacity to adapt to adverse
conditions during transplanting and establishment under
harsh environments (Table I). After transplantation, and
throughout the establishment period, pre-conditioned
plants show higher l, p, and RWC values, lower os

Main morphological and physiological characteristics of seedlings at transplantation and nursery pre-conditioning techniques (microclimate and cultivation
methods) which improve the establishment and survival of ornamental plants under harsh environments in xerogardening and landscaping
Seedling Characteristic
Morphological
Low shoot:root ratio,
low height and leaf area
Thick roots
Metacutisated rootsy
Mycorrhizal symbiosis
TABLE I
Nursery Technique Species Reference
Deficit irrigation Acer pseudoplatanus Hipps et al. (1996)
Limonium cossonianum Franco et al. (2002b)
Lotus creticus Franco et al. (2001; 2002a)
Myrtus communis Bañón et al. (2002)
Rosmarinus officinalis Sánchez-Blanco et al. (2004b)
Low air temperature L. creticus Bañón et al. (2004)
Picea glauca, Pinus contorta Van Den Driessche (1991a)
Low air humidity M. communis, N. oleander Bañón et al. (2002; 2005b)
Enriched CO2 air Pinus nigra Biel et al. (2004)
Paclobutrazol Arbutus unedo Navarro et al. (2004)
Phillyrea angustifolia Fernández et al. (2004)
Pyracantha sp. Ruter (1994)
Root pruning Conifer Aldrete and Mexal (2002)
Larger container sizes Ornamental shrubs Pastor et al. (1999)
Light-weight substrate Viburnum tinus Pastor et al. (1999)
Thuja occidentalis Hinesley et al. (1999)
Moderate N supplies Picea abies Rikala et al. (2004)
Deficit irrigation M. communis, N. oleander Bañón et al. (2002; 2005b)
Deficit irrigation L. cossonianum, L. creticus Franco et al. (2001; 2002b)
Mycorrhizal inoculation Ornamental plants Carpio et al. (2003)
R. officinalis Sánchez-Blanco et al. (2004a)

Physiological
Low gs, l, p and RWCz Deficit irrigation L. creticus Bañón et al. (2004)
M. communis, N. oleander Bañón et al. (2002; 2005b)
R. officinalis Sánchez-Blanco et al. (2004b)
Low air humidity M. communis, N. oleander Bañón et al. (2002; 2005b)
Olea europaea var. sylvestris Bañón et al. (2003b)
Pistacia lentiscus Vilagrosa et al. (2003)
Rhamnus alaternus Bañón et al. (2003c)
R. officinalis Sánchez-Blanco et al. (2004b)
Low air temperature Eucalyptus spp. Close et al. (2004)
P. glauca, P. contorta Van Den Driessche (1991b)
y
Process of lignification and suberisation that results in a resting root that is protected against fluctuations in environmental conditions and is capable
of re-growth when conditions ameliorate.
gs, stomatal conductance; l, leaf water potential; p, leaf turgor potential; RWC, relative water content.
z
 J. A. FRANCO, J. J. MARTÍNEZ-SÁNCHEZ, J. A. FERNÁNDEZ and S. BAÑÓN
values, and maintain their gs, whereas a substantial fall in
gs is observed in non-pre-conditioned plants.
Selecting ornamental plants for improved nursery
performance and tolerance of harsh environments has
long been of interest to those involved in landscape
horticulture, urban greenspace design and xerogardening
projects. The use of native species of wild flora is of
increasing interest because of their capacity to adapt to
adverse environmental conditions (Prevete et al., 2000;
Savé et al., 2000; Franco et al., 2002b; Torrecillas et al.,
2003; Reaves and Whitlow, 2004; Sánchez-Blanco et al.,
2004c; Sharma and Graves, 2004; Schrader and Graves,
2004; Shoemake et al., 2004). However, it is known that
the degree of adaptation to abiotic stresses may vary
considerably within a family, within a genus and even
within a species.
Several factors and techniques influence seedling
production and performance (Table I). In general,
seedlings growing under very high or very low moisture
conditions are adversely affected, while those growing
under moderate conditions exhibit optimum growth, bud
development and have sufficient nutrient and starch
reserves. The restricted growth observed in the nursery
under low temperatures, low air humidity and, mainly,
deficit-irrigation, can be considered a morphological
adaptation of the plant to water and environmental
stresses to reduce transpiration and lower the
consumption of water. Stomatal density also plays an
important role in the response of plants to hardening
during the nursery phase. However, stomatal regulation
may also be relevant as a mechanism to reduce
transpiration and water loss through the leaves (Van
Den Driessche, 1991a, b; Sánchez-Blanco et al., 2002;
Vignolio et al., 2002; Clary et al., 2004; Bañón et al.,
2005b).
The root system is as important, or even more
important, than the aerial parts of the plant for successful
transplanting and establishment in the field. However,
the influence of environmental conditions and
cultivation techniques in the nursery on the root
characteristics of seedlings and on post-transplant root
development has been little studied. Undoubtedly, this is
due to limitations on accessibility for root observations.
Studies on root system dynamics are especially difficult
because they require successive, non-destructive
measurements. Having an optimum root system at
transplantation and the production of primary roots may
be decisive features for plant survival. Deficit-irrigation
during the nursery phase can increase root-length and
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13
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