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Why seedlings survive: influence of plant attributes
Steven C. Grossnickle
Received: 25 October 2011 / Accepted: 25 April 2012 / Published online: 13 May 2012
Springer Science+Business Media B.V. 2012
Abstract Seedling survival and successful forest restoration involves many silvicultural
practices. One important aspect of a successful forest restoration program is planting
quality seedlings with high survival capability. Thus the nursery needs to create seedlings
with plant attributes that allow for the best chance of success once a seedling is field
planted. Since the mid-twentieth century, research foresters have critically examined plant
attributes that confer improved seedling survival to field site conditions. This review
describes the value of commonly measured seedling quality material (i.e. shoot height,
stem diameter, root mass, shoot to root ratio, drought resistance, mineral nutrient status)
and performance (i.e. freezing tolerance and root growth) plant attributes defined as
important in answering the question of why seedlings survive after planting. Desirable
levels of these plant attributes can increase the speed with which seedlings overcome
planting stress, become ‘coupled’ to the forest restoration site, thereby ensuring successful
seedling establishment. Although planting seedlings with these desirable plant attributes
does not guarantee high survival rates; planting seedlings with desirable plant attributes
increases chances for survival after field planting.
Keywords Seedling survival Forest restoration Morphological attributes Drought
resistance Freezing tolerance Seedling nutrition Root growth capability
Introduction
Why seedlings survive after planting has long been debated because seedling survival is
pivotal in the initial success of a forest restoration program. During the early part of the
twentieth century, programs planting nursery-grown seedlings in North America reached
an annual size of 10–20 million (Toumey 1916). Due to this silvicultural investment,
foresters began examining plantation failures and tried to discern reasons for seedling
mortality (e.g. Tillotson 1915; Young 1921; Kittredge 1929; Rudolf 1939). Often seedling
S. C. Grossnickle (&)
CellFor Inc., Saanichton, Canada
e-mail: sgrossnickle@shaw.ca
123
New Forests (2012) 43:711–738
DOI 10.1007/s11056-012-9336-6
losses were attributed to environmental stress, animal grazing, disease, or insects. How-
ever, in many cases poor quality planting stock (Kittredge 1929) or the inability of planted
seedlings to grow roots (Rudolf 1939) was defined as the cause of plantation failure. The
work of Toumey (1916) and studies initiated on southern pines in the 1920s (reported by
Wakeley 1954) were some of the initial attempts to grade nursery stock using morpho-
logical parameters to improve seedling establishment. Thus, early in the twentieth century
researchers began to ask the question of how plant attributes influence seedling survival
after field planting.
Nursery cultural and silvicultural practices have a strong influence on seedling per-
formance immediately after planting. Effects of these practices on seedling performance
need to be understood to make sound forest restoration decisions. Regeneration silviculture
is a complex process and many factors go into making a successful forest restoration
program. Implicit within a seedling production program is recognition of the inherent
species characteristics when making the proper selection of the genetic source that is
adapted to forest restoration site conditions (Zobel and Talbert 1984). Also silvicultural
practices related to the regeneration process (e.g. storage, handling, planting date, planting
practices, site preparation, and vegetation management) all have an effect on the success of
a forest restoration program. Readers interested in understanding effects of these factors
can examine a number of excellent texts on silvicultural practices and forest regeneration
site performance of planted seedlings (e.g. Cleary et al. 1978a; Lavender et al. 1990;
Duryea and Dougherty 1991; Hobbs et al. 1992; Grossnickle 2000; Wagner and Colombo
2001).
How recently planted seedlings initiate growth and become ‘‘coupled’’ into the forest
ecosystem (Grossnickle 2005a), thereby avoiding water stress, are critical factors for
success of a forest restoration program. It is the lack of coupling (i.e. due to a restricted
rooting volume limiting access to soil water) that increases the possibility of excessive
water stress in seedlings which can result in either carbon starvation or hydraulic failure,
and subsequently seedling death (McDowell et al. 2008). Thus, seedling survival is related
to their inherent growth potential and the degree to which field site environmental con-
ditions limits or enhances this potential for seedlings to become established or coupled into
the forest ecosystem (Grossnickle 2000).
In the mid-twentieth century, researchers started to critically examine plant attributes
that conferred improved survival for bareroot (Wakeley 1948,1954; Stone 1955) and
container-grown (Tinus 1974) seedlings. This was the start of seedling quality programs
based on the need for a better understanding of performance capabilities of nursery-grown
seedlings in relation to the forest restoration site. Seedling quality assessment has evolved
to include numerous morphological and physiological measurement procedures for
defining field performance (Mattsson 1996; Grossnickle 2000). Subsequently, a wealth of
information has been published on plant attributes that improve the odds of survival once a
seedling is field planted.
Defining seedling quality comes from measurements of seedling properties that describe
material (i.e. single point measures of individual plant parameters) and performance (i.e.
plant measurements reflecting an integrated response of many material attributes to defined
environmental conditions) attributes (Ritchie 1984). This review examines the seedling
through commonly measured material (i.e. shoot height, stem diameter, root mass, shoot to
root ratio, drought resistance, mineral nutrient status) and performance (i.e. freezing tol-
erance and root growth) plant attributes used to define seedling quality. The objective of
this review is to conduct a comprehensive, though not exhaustive, examination of work
712 New Forests (2012) 43:711–738
123
describing the value of primary plant attributes that are important in answering the question
of why seedlings survive after planting on forest restoration sites.
Material morphological attributes
…serve their purpose only so far as they actually separate seedlings with a high
capacity for survival and growth after planting from those with a low capacity.
(Wakeley 1948)
Extensive work since the 1950s shows desirable morphological attributes contribute to
seedling survival after transplanting on to forest restoration sites. To summarize this work,
a well-balanced shoot to root system, with a sturdy stem and a large fibrous root system
provides the best chance for seedling survival (e.g. southern pines—Lantz 1985; South
2000, radiata pine (Pinus radiata D. Don)—Menzies et al. 1985, Pacific Northwest and
Northern British Columbia tree species—Scagel et al. 1993, temperate zone deciduous
hardwoods—Wilson and Jacobs 2006). Morphological attributes are considered a reliable
measure of seedling quality (Puttonen 1997) because they retain their mark on the seedling
identity for extended timeframes after seedlings are field planted and start to grow. These
plant structural features play a key role in defining their hydraulic architecture (i.e.
potential water balance), thus helping to determine whether plants live or die during
exposure to drought (McDowell et al. 2008).
Even so, historical work has found morphological attributes to be an inconsistent
measure of seedling survival (Wakeley 1954; Thompson 1985; Mexal and Landis 1990).
This stems from the fact that morphological parameters only measure overall seedling size,
growth potential and shoot to root balance, not seedling physiological quality. Thus
measures of morphological parameters are only part of the equation of plant attributes
required for successful seedling survival (Wakeley 1948,1954; Tinus 1974; Ritchie 1984;
Mexal and Landis 1990). With this caveat in mind, the following discussion focuses on the
influence morphological parameters have on seedling survival.
Shoot height
Large seedlings have been recommended for planting on sites where there is little envi-
ronmental stress but there is the potential for excessive competition (Toumey 1916). As
black spruce (Picea mariana (Mill.) B.S.P.) seedlings increased in shoot size they had a
greater exposure to growing season available light on sites with high competition, resulting
in greater survival and shoot growth (Jobidon et al. 1997,2003). Taller bareroot loblolly
pine (Pinus taeda L.) seedlings at planting confer higher survival on sites with little
environmental stress (Fig. 1). Numerous studies show large stock on sites where compe-
tition is prevalent can improve survival and growth (Newton et al. 1993; Mason et al. 1996;
Mohammed et al. 1998; South and Mitchell 1999; Pue
´rtolas et al. 2003; Villar-Salvador
et al. 2004a).
Shoot height is a general measure of photosynthetic and transpirational capacity and
also a reflection of potential height growth (Armson and Sadreika 1979; Cleary et al.
1978b; Mexal and Landis 1990). As the shoot system initiates growth, larger seedlings
produce greater absolute amounts of new shoot biomass (Thiffault 2004; Grossnickle
2005b) and occupy a greater area within the planting spot than smaller seedlings, thereby
capturing more incoming solar radiation. This is critical because the effect of competition
New Forests (2012) 43:711–738 713
123
on limiting sunlight, thus seedlings net photosynthetic carbon gain, is related to survival
(Johnson and Smith 2005). Shoot system size is important because on sites with available
soil water and nutrients, competition for light between planted seedlings and the site
vegetation complex is a main factor limiting seedling performance (Grossnickle 2000).
In contrast, planting seedlings with greater height can result in lower survival on dro-
ughty sites (Larsen et al. 1986; Boyer and South 1987; Tuttle et al. 1988; McTague and
Tinus 1996). For example, shorter bareroot loblolly pine seedlings had higher survival on
sites with limited soil water and greater environmental stress (Fig. 1). Under dry soil
conditions, larger, compared to smaller, conifer seedlings can have greater water stress
(Rose et al. 1993; Stewart and Bernier 1995), lower photosynthesis (Lamhamedi et al.
1997) and reduced growth (Baer et al. 1977; Hahn and Smith 1983). As the shoot system
reaches a certain size, increased foliar mass can increase the seedling’s susceptibility to
planting stress because a newly planted seedling’s root system cannot supply enough water
to transpiring foliage to maintain a proper water balance (Grossnickle 2005a). The sus-
ceptibility of larger seedlings being exposed to water stress at planting is mitigated if they
have the capability to quickly develop new roots (see Root Growth).
Stem diameter and root mass
Seedling stem diameter is a general measure of seedling sturdiness, root system size, and
protection against drought and heat damage (Cleary et al. 1978b; Mexal and Landis 1990).
Thus, it is difficult to separate the relationship between these two parameters, with a greater
root system size occurring as stem diameter increases in both bareroot (Ritchie 1984) and
container-grown (Grossnickle 2000) seedlings. Greater root mass is an indicator of root
absorptive surface (Thompson 1985) conferring a greater seedling drought avoidance
capability after field planting.
Johnson and Cline (1991) considered stem diameter the single most useful morpho-
logical measure of seedling quality. Numerous studies show larger stem diameter seedlings
tend to survive better than small stem diameter seedlings (e.g. bareroot; van den Driessche
1980,1984; South and Mexal 1984; Long and Carrier 1993; McGrath and Duryea 1994;
South 1993; Zwolinski et al. 1996; South and Mitchell 1999; South et al. 2001,2005; Rose
and Ketchum 2003; Morrissey et al. 2010) (e.g. container-grown; Hines and Long 1986;
Fig. 1 The relationship between survival (year 2) and initial seedling height for bareroot loblolly pine
(Pinus taeda L.) seedlings on adverse and non-adverse sites (Tuttle et al. 1987)
714 New Forests (2012) 43:711–738
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Bayley and Kietzka 1997; Mexal et al. 2008; South et al. 2005; Oliet et al. 2009b;
Morrissey et al. 2010). This is why Mexal and Landis (1990) felt stem diameter can best
forecast field survival because it indirectly confers a number of desirable plant attributes
(i.e. water absorption—roots, water transport—stem) that are considered important
parameters of the plant hydraulic architecture that play a role in plant survival during
drought (McDowell et al. 2008).
The relationship of larger seedling stem diameter, and by extension root development,
and survival is not universal. In a review of bareroot pine seedlings spanning many trial
sites in the southeastern US, survival was related to initial stem diameter at planting in
most, but not all cases (Fig. 2). As seedling stem diameter increased in size the probability
that a plantation would have survival at\75 % declined from a high of 88 % for seedlings
with a small stem diameter (i.e.\2.4 mm) to only 9 % for seedlings with a very large stem
diameter (i.e.[6.3 mm). Seedlings with a larger stem diameter had a greater probability of
high survival, but not in all field situations.
Seedlings with more roots have better survival potential (Toumey 1916) and seedlings
with a good quality root system have better establishment capability after field planting
(Davis and Jacobs 2005). Greater root system size at planting can result in greater seedling
survival (Larsen et al. 1986; Rose et al. 1997). Seedlings with a larger stem diameter and
more roots exhibit fewer symptoms of planting stress (Haase and Rose 1993) and have
better survival than seedlings with a smaller stem diameter and fewer roots (South et al.
1985; Carlson 1986; Blake et al. 1989; Hobbs et al. 1989; Long and Carrier 1993). Greater
root system size can confer greater root growth capability (see Root Growth). On harsh
sites, a greater capability for absorption and transport of water from roots through the stem
to the transpiring shoot system gives seedlings a better chance of overcoming planting
stress (Grossnickle 2005a).
Shoot to root ratio
Measures of shoot to root (S:R) balance define seedlings drought avoidance potential
(Thompson 1985). The need for seedlings to have root systems in proper proportion to the
Fig. 2 The probability of seedling survival in the field \75 % for bareroot loblolly pine (Pinus taeda L.)
and slash pine (Pinus elliottii Engelm.) seedlings graded into stem diameter classes (adapted from South
et al. 1985). Note: Number within a bar for a stem diameter class defines the number of field sites from
where data was collected
New Forests (2012) 43:711–738 715
123
shoot system has long been recognized as a desirable seedling attribute (Toumey 1916)
because seedling water status is directly tied to their S:R (Parker 1949). An imbalance in
the shoot transpirational surface to the root absorbing surface (i.e. S:R [3.0) can result in
water stress for bareroot seedlings (Baldwin and Barney 1976). A S:R between 1.0 and 3.0
gave bareroot seedlings a better chance of survival (Hermann 1964; Foiles and Curtis
1973; Hobbs 1982), with survival increasing as S:R decreases (Tanaka et al. 1976;
Lopushinsky and Beebe 1976; Larsen et al. 1986; Boyer and South 1987; Chamshama and
Hall 1987; Kainer and Duryea 1990; Haase and Rose 1993;Ge
´ne
´re
´and Garriou 1999)
(Fig. 3a). Thompson (1985) found bareroot seedlings with lower S:R on average had a
29 % higher level of survival. The bareroot nursery cultural practices of undercutting
(reviewed by South and Donald 2002) and top pruning (South and Blake 1994) are
commonly used to decrease the S:R and produce seedlings with better survival capability.
Even though data showing the importance of S:R for bareroot seedling survival is com-
pelling, Hobbs (1984) warns that assessing potential survival based on the S:R without also
judging the quality of the root system (i.e. fibrous root system with many growing tips) can
limit its reliability to forecast survival.
Fig. 3 a The effect of shoot to root ratio of bareroot loblolly pine (Pinus taeda L.) seedlings on the survival
under a simulated drought environment (adapted from Mexal and Dougherty 1983). bThe effect of shoot to
root ratio of container-grown Patula pine (Pinus patula Schiede ex Schltdl. & Cham.) seedlings on survival
1 month after field planting (adapted from Bayley and Kietzka 1997)
716 New Forests (2012) 43:711–738
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Studies have also found the survival of container-grown seedlings to be greater with
lower S:R under droughty field conditions (Zida et al. 2008; del Campo et al. 2010)
(Fig. 3b). While these reports show the importance of initial S:R to enhance survival for
container-grown seedlings, S:R based on total shoot and root weights may be limited in
forecasting survival except under harsh field conditions. Bernier et al. (1995a) found little
evidence that the S:R based on total shoot and root weights forecast survival for container-
grown seedlings on sites lacking severe environmental stress. Bernier et al. (1995a) argued
that the root plug-soil interface is the primary limiting factor rather than the overall root
system in affecting seedling performance. Container-grown seedlings are typically grown
in a medium that has desirable characteristics for root growth in the nursery (i.e. increased
aeration and water holding capacity, low bulk density) (Tinus and McDonald 1979). These
media characteristics limit movement of water into the root plug, after field planting, due to
physical constraints of low density peat plugs (Bernier 1992; Bernier et al. 1995b) causing
an imbalance of soil matric potential between plug media and field site soil (Day and
Skoupy 1971). The ratio of root development outside of the container plug to total shoot
dry weight best represented container-grown seedling drought avoidance potential; seed-
lings with minimal root development outside the plug having the greatest level of water
stress (Grossnickle and Reid 1984). This is because the root plug-soil interface is the point
of greatest resistance to water flow for newly planted container-grown seedlings (O
¨rlander
and Due 1986). A measure of root development out of the plug and into the soil in relation
to shoot mass is a logical description of S:R balance of container-grown seedlings that
better reflects their drought avoidance, thus survival potential.
Material and performance physiological attributes
…seedlings must be produced in such a way as to be physiologically ready to
outplant into the field environment. (Lavender and Cleary 1974)
Survival is determined, in part, by the ability of seedlings to respond to environmentally
stressful conditions that can occur after being planted. The most dramatic of these site
conditions are alterations in the heat exchange processes and site-water relations limiting
seedling performance (Miller 1983), while site nutrient dynamics are also considered rate
limiting to young forest stands (Troth et al. 1986). The following discussion focuses on
physiological attributes that allow seedlings to survive conditions of drought, frost and
nutrient dynamics just after planting on a forest restoration site.
Cultural practices that provide an improved ‘‘physiological quality’’ to seedlings have
long been considered important in increasing their chances for survival just after field
planting (Wakeley 1948,1954). This is because nonhardened seedlings (Rowe 1964; Tinus
1974; Hobbs 1984) or seedlings lacking the proper nutrient balance (see Nutrient Status)
lack the physiological capability to become rapidly established after planting on forest
restoration sites. Acclimation of seedlings is based on the concept of ‘‘slowly increasing
stresses to induce physiological adjustments in plants’’ (Kozlowski and Pallardy 2002)
thereby developing protection from potentially stressful field site conditions. Thus, plant
acclimation or nursery cultural hardening practices (Wakeley 1954; Lavender and Cleary
1974; Landis et al. 1999) are applied to increase the odds in favor of seedling survival to
field site conditions. Nursery practitioners have used cultural practices of reduced day-
length, temperature, watering and fertilization regime modification to harden container-
grown seedlings (Tinus and McDonald 1979; Landis et al. 1999), with watering and
New Forests (2012) 43:711–738 717
123
fertilization regime, plus shoot and root culturing modification used to harden (also
improve S:R) bareroot seedlings (Duryea 1984; Mexal and South 1991). These treaties
provide a detailed explanation of cultural practices used to harden seedlings. The following
sections discuss physiological attributes, adjusted by hardening practices, which can
improve seedlings chances of survival after transplanting to forest restoration sites.
Drought resistance
Drought stress causes tree species to develop drought resistance (Abrams 1988). Nurseries
apply cultural practices that create water stress events at the end of the growing season to
trigger budset or the cessation of shoot growth, and initiate stress resistance in seedlings.
Periodic moderate water stress is used as a nursery cultural practice to induce bud for-
mation (Lavender and Cleary 1974; Timmis and Tanaka 1976; Young and Hanover 1978;
Macey and Arnott 1986; Calme
´et al. 1993) as well as improve drought resistance (i.e.
tolerance and avoidance) through osmotic adjustment (Kandiko et al. 1980; Hennessey and
Dougherty 1984; Buxton et al. 1985; Ritchie and Roden 1985; Seiler and Johnson 1985;
Grossnickle et al. 1991a; Major et al. 1994; Villar-Salvador et al. 2004b), stomatal sen-
sitivity to drought (Unterschuetz et al. 1974; Roberts and Dumbroff 1986; Zwiazek and
Blake 1989; Villar-Salvador et al. 1999), reduce susceptibility to xylem cavitation in some
species (Beikircher et al. 2010), but not all species (Harvey and van den Driessche 1999;
Beikircher et al. 2010), and create seedlings with lower S:R (Timmer and Miller 1991;
Bayley and Kietzka 1997; Biel et al. 2004; Thomas 2009; Verdauger et al. 2011).
Adjustment of the fertilization regime (i.e. from a high to lower N concentration plus
improving the P and K status) in combination with drought hardening can improve drought
resistance in hybrid poplars (Populus spp.) (Harvey and van den Driessche 1997,1999).
Short-day treatments that initiate budset can enhance drought resistance in the summer
(Grossnickle and Folk 2003; Tan 2007) and fall (Colombo 1987; Grossnickle et al. 1991a;
Major et al. 1994) crops of temperate zone tree species, though an extended short-day
regime can be counterproductive causing increased mortality of summer-planted white
spruce (Picea glauca (Moench) Voss) seedlings (Tan et al. 2008).
The use of hardening practices can improve the performance of field planted seedlings
(Kozlowski et al. 1991). For example, drought hardening allowed seedlings to have a faster
recovery of photosynthetic capability after transplanting (Kaushal and Aussenec 1989),
better control of water loss (Christersson 1972; Clemens and Jones 1978; Johnson et al.
1985; Timmer and Miller 1991; Villar-Salvador et al. 1999,2004b), and greater root
regeneration after planting into soils having limiting edaphic conditions (i.e. droughted
soils—Kaushal and Aussenec 1989, and cold soils—Hennessey and Dougherty 1984;
Arnott et al. 1994). This improved performance has, in some cases, resulted in drought
hardened seedlings having increased survival (Blake et al. 1979; van den Driessche 1991a,
1992) (Fig. 4) and growth when planted in xeric soil conditions (Johnson et al. 1985).
Drought hardening cultural practices does not always have beneficial effects on seedling
field survival. This cultural practice can have either no effect (Grossnickle et al. 1991b;
Folk et al. 1994; Royo et al. 2001; Biel et al. 2004; Villar-Salvador et al. 2004b)ora
negative effect (Jospon and Paul 1985; O’Reilly et al. 1994) on survival in the field. This
discrepancy probably stems from stress intensity during nursery application (Kozlowski
and Pallardy 2002), timing of hardening practices (Duryea and McClain 1984; Landis et al.
1999), or the fact that stress resistance is not required to ensure seedling survival when
exposed to optimum field conditions.
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Freezing tolerance
Drought hardening practices can improve freezing tolerance in some conifer species
(Timmis and Tanaka 1976; Blake et al. 1979; Grossnickle et al. 1991a). However, this
phenomenon has not been observed in other conifers (van den Driessche 1969; D’Aoust
and Cameron 1982; Menzies et al. 1981; Arnott et al. 1994). Drought hardening can
improve seedling survival after field planting into frost prone conditions (Mexal et al.
1979).
Most tree species undergo many morphological and physiological changes during an
annual phenological cycle in response to seasonal environmental conditions (Fuchigami
and Nee 1987; Ritchie and Tanaka 1990; Burr 1990) with freezing tolerance (Burr 1990;
Bigras et al. 2001) shifting to its highest level in the winter in many tree species. This
phenomenon has been related to budset or the cessation of shoot growth, leaf maturation
and seasonal shifts in temperature (Grossnickle 2000). Short-day treatments are an
effective means to initiate budset (Vaartaja 1960; Lavender and Wareing 1972; Williams
et al. 1972), control dormancy patterns (Lavender and Cleary 1974; Lavender 1985) and
enhance freezing tolerance in the fall (Colombo et al. 2001) for temperate zone tree
species. Proper fall hardening of temperate zone tree species ensures they can be lifted and
stored for extended periods to maintain a high level of seedling quality (Colombo et al.
2001) without a depletion of carbohydrate reserves below a critical level that can affect
survival (Ritchie 1982; Marshall 1985). These practices result in increased seedling sur-
vival (McKay and Mason 1991) (Fig. 5), improved capability to overcome planting stress
(Grossnickle 2000) and become established when planted into the spring planting window
(Grossnickle et al. 1991a,b,c; Jacobs et al. 2008).
However, short-day treated spruce (Picea spp.) seedlings can show earlier budbreak
setting up the potential for damage of actively growing seedlings by early-spring frost
(Grossnickle 2000). Terminal bud damage of short-day treated spruce seedlings is attrib-
uted to early growing season frost, although this only occurs on frost-prone sites (Kra-
sowski et al. 1993). This is a possible limitation of this cultural practice to maintain desired
Fig. 4 Drought avoidance response of control and drought hardened (mild and severe drought (DR)
culture) Eucalyptus pilularis Sm. seedlings to drought conditions in a glasshouse (adapted from Thomas
2009) (Insert data shows the shoot to root ratio of seedlings after drought hardening cultural conditions with
different letters indicating treatment differences (p=0.05))
New Forests (2012) 43:711–738 719
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freezing tolerance levels, thereby limiting damage or mortality when planting seedlings
into frost prone conditions.
Nutrient status
Proper nutrient balance
During the nursery acclimation process, fertilization practices are shifted in concert with
other cultural parameters to slow and then cause growth cessation in container-grown
(Landis et al. 1989) and bareroot (Duryea 1984; Lantz 1985) seedlings. This approach was
developed under long held fertilization adjustment practices (bareroot seedlings—Wakeley
1954, container-grown seedlings—Lavender and Cleary 1974) to ensure seedlings had
budset or the cessation of shoot growth at the proper time to develop hardiness and still go
to the field with sufficient nutrient levels for good survival and growth. Sufficient nutrient
levels at planting are critical because seedlings have a limited ability to access required
nutrients from the field site during the establishment process (Tinus 1974).
Nursery fertilization can affect seedling survival after planting. In a review of 22 trials
van den Driessche (1991b) found that ‘‘suitable nursery fertilization’’ resulted in 57 % of
trials showing an increase in survival, 30 % of trials showing no effect and 13 % reported a
decrease in seedling survival in the field. In a series of trials, survival of Douglas-fir
(Pseudotsuga menziesii (Mirb.) Franco) seedlings was related to N concentration at
planting, with a 2 % (±0.5 %) N concentration resulting in the highest rate of survival 1
and 2 (Fig. 6), and 3 years (van den Driessche 1984,1988) after planting.
Nutrient loading
The shift of fertilization practices after budset or the cessation of shoot growth to increase
nutrient concentration (i.e. fall nutrient loading) with little effect on seedling development
has long been considered a beneficial nursery cultural practice (Benzian et al. 1974; Brix
and van den Driessche 1974). Fall fertilization after seedlings have ceased growth, can
Fig. 5 Relationship between first year field survival and pre-storage freezing tolerance (measured as the
temperature at which 50 % needle electrolyte leakage occurred: LT
50
) for lodgepole pine (Pinus contorta
Dougl.) seedlings (Simpson 1990)
720 New Forests (2012) 43:711–738
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increase the nutrient status without delaying progression of the bud dormancy cycle
(Williams and South 1992), while not affecting (Luoranen et al. 2008) or even improving
(Islam et al. 2009; Andivia et al. 2012) the fall development of freezing tolerance. Ingestad
and Lund (1986) theorized that nutrient loading in the nursery provided seedlings with
greater nutrient reserves to utilize after field planting, while Binkley (1986) considered
increasing nutrient reserves through nursery fertilization a very energy-efficient approach
to nutrient acquisition for recently planted seedlings, compared to the uptake of nutrients
from the soil. Fall nutrient loading can also contribute to enhanced stress resistance
(Timmer 1997), shoot growth potential (i.e. increased needle primordial in buds) (Colombo
et al. 2003; Islam et al. 2009) and a lower S:R at planting (Timmer et al. 1991). Fall
nutrient loading increases the availability of nutrient reserves that are rapidly remobilized
to support nutrient demand of new growth once seedlings are planted.
Application of fall nutrient loading has a number of effects on the physiological
response of seedlings after field planting. Reports show fall nutrient loading results in
earlier shoot flush (Margolis and Waring 1986; Floistad and Kohmann 2004; Oliet et al.
2011), increased new root growth (van den Driessche 1985,1988,1992; Malik and
Timmer 1996,1998; Boivin et al. 2004; Villar-Salvador et al. 2004a; Oliet et al. 2009a,
2011; Andivia et al. 2011,2012), increased nutrient uptake (Timmer and Aidelbaum 1996)
and shoot growth (van den Driessche 1985,1992; Margolis and Waring 1986; Malik and
Timmer 1996; VanderSchaaf and McNabb 2004; Pue
´rtolas et al. 2003; Boivin et al. 2004;
Close et al. 2005; Salifu et al. 2009; Oliet et al. 2009a,2011). Nutrient loaded seedlings can
have a greater capability to overcome planting stress on harsh sites and they can be an
effective stocktype on forest restoration sites with high levels of competition (van den
Driessche 1991b; Timmer 1997).
In some instances, fall nutrient loading improves seedling survival in the field (van den
Driessche 1980,1984,1992; Irwin et al. 1998; Oliet et al. 2009b; del Campo et al. 2010),
while in other field trials minimal survival benefits were noted (Benzian et al. 1974; van
den Driessche 1988; Gleason et al. 1990; Birchler et al. 2001; South and Donald 2002;
VanderSchaaf and McNabb 2004; Salifu et al. 2009; Andivia et al. 2011). In a few
instances nutrient loading reduced seedling survival in the field (Benzian et al. 1974; South
and Donald 2002; Boivin et al. 2004) and Boivin et al. (2004) attributed this phenomenon
to excessive nutrient loading causing toxic nutrient concentrations. Variability in the
Fig. 6 Seedling survival at 1 and 2 years after field planting in relation nitrogen concentration for Douglas-
fir (Pseudotsuga menziesii (Mirb.) Franco)) seedlings at planting (van den Driessche 1980)
New Forests (2012) 43:711–738 721
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response to nutrient loading indicates that this practice may only be beneficial when site
nutrient restrictions are limiting seedling establishment.
Seedlings loaded with nutrient reserves can have unintended consequences to forest
restoration programs. Higher mortality for nursery-grown spruce seedlings by grazing from
snowshoe hares (Lepus americanus Erxleben) was attributed to preferential feeding on
nursery-grown than naturally regenerated seedlings (Sullivan and Moses 1986; Rodgers
et al. 1993). A similar phenomenon of herbivores feeding occurs in conifers (Bergquist and
O
¨rlander 1998; Burney and Jacobs 2011) and hardwoods (Close et al. 2004; Paul et al.
2012) where the frequency of browsing by herbivores was greater with higher N con-
centration. However, increased browsing was not always associated with N levels (Burney
and Jacobs 2011; Paul et al. 2012) and on sites with high browse pressure seedlings with
both high and low N levels were browsed (Close et al. 2004). Increasing nutrient reserves,
through nursery fertilization, may decrease seedling field survival due to browsing damage
of herbivorous animals.
Shift in physiological attributes
Since effects of drought and cold hardening can be tied to a plants normal phenological
cycle, these hardening benefits are ephemeral in nature. As tree species initiate shoot
growth in the spring, drought tolerance (Teskey and Hinckley 1986; Abrams 1988;
Grossnickle 2000) and freezing tolerance (Burr 1990; Bigras et al. 2001) can be lost in
rapid fashion. For example, interior spruce (Picea glauca (Moench) Voss 9Picea eng-
elmannii Parry) seedlings lost a good portion of their stress resistance within weeks of
initiating growth (Fig. 7).
Any potential benefit of increased fertility in the nursery in terms of improved seedling
performance in the field is also short-lived. Seedling nutrient reserves decline after
planting, due to dilution in tissue nutrient concentrations if external nutrient sources cannot
Fig. 7 Shift in physiological attribute status of nutrient loaded spring planted black spruce (Picea mariana
(Mill.) B.S.P.) seedlings (NUT Load; adapted from Malik and Timmer 1998), and drought tolerance (DR
TOL—osmotic potential at turgor loss point), drought avoidance (DR AVD—cuticular transpiration) and
freezing tolerance (FR TOL—Index of Injury at -6C), of spring planted interior spruce (Picea glauca
(Moench) Voss 9Picea engelmannii Parry) seedlings (adapted from Grossnickle and Folk 2007) at time of
planting and 5–7 weeks after planting. Note: Seedlings had budbreak 2–3 weeks after planting
722 New Forests (2012) 43:711–738
123
meet demands of new growth (Munson and Bernier 1993; Kim et al. 1999). Nutrient
loaded black spruce seedlings lost 26 % of their N concentration 7 weeks after they
resumed growth (Fig. 7) and N concentration of nutrient loaded and control seedlings were
comparable by the end of the growing season (Malik and Timmer 1998).
Seedlings ability to utilize improved physiological plant attributes to overcome planting
stress and become established is a very narrow window, making it very difficult to quantify
benefits of these hardening and nutrient loading practices on survival. For this reason,
seedling survival and successful establishment is not only predicated on their hardiness and
nutrient status, but also on their morphological attributes (see above discussion) and
capability to grow roots (see below discussion) after planting.
Performance attribute: root growth
If the root system did not increase in size at a fairly rapid rate…the seedling would
die of drought…. (Stone 1955)
Seedlings that develop a root system after planting establish a proper water balance
because they are coupled into the hydrologic cycle whereby water flows from the soil to
plant roots, through the plant and into the atmosphere (Grossnickle 2005a). If sufficient
root development does not occur just after field planting, seedlings can be exposed to stress
because they do not have access to soil water. This planting stress can lead to a cycle of
root growth being limited by the lack of water and photosynthates, and in turn photo-
synthesis being limited by water stress due to a lack of root growth (Burdett 1990;
Grossnickle 2000). Alternatively, seedlings that establish roots quickly after planting
develop a favorable water status which continues the cycle of root growth supported by
photosynthesis (Guehl et al. 1989) and photosynthesis supported by root growth (Burdett
1990). This is why survival is predicated on sufficient root growth coupling the newly
planted seedling to the site, thereby maintaining a proper plant water balance (Margolis
and Brand 1990; Grossnickle 2005a).
This view that root growth is critical for seedling survival is why root growth capability
is reported to be a common measurement tool used in operational programs worldwide to
define seedling quality (Simpson and Ritchie 1997). This assessment approach is deter-
mined through a testing procedure called root growth capacity or root growth potential.
Numerous reviews have discussed merits of measuring root growth within a seedling
quality assessment approach for determining seedling performance (Ritchie and Dunlap
1980; Ritchie 1985; Burdett 1987; Ritchie and Tanaka 1990; Sutton 1990). This assess-
ment approach is considered a direct indicator of a seedlings ability to grow roots and is a
general indicator that all physiological systems are functioning properly and thus provides
a measure of seedling performance potential (Ritchie 1984; Burdett 1987). This is why root
growth in newly planted seedlings has long been recognized as important to ensure suc-
cessful survival and establishment (Toumey 1916; Rudolf 1939; Wakeley 1948,1954;
Stone 1955; Tinus 1974).
Due to the necessity of root growth in successful seedling establishment, a critical
aspect of nursery cultural programs is to produce seedlings with the capability to rapidly
grow roots after field planting. Factors of seedling size, root system fibrosity and stocktype
affect their capability to grow roots.
New Forests (2012) 43:711–738 723
123
Seedling size
Seedling size affects a seedling’s ability to grow roots. Greater initial root mass is related
to greater root growth (Brissette and Roberts 1984; Johnsen et al. 1988; Williams et al.
1988; van den Driessche 1992; Grossnickle and Major 1994; Villar-Salvador et al. 2004a;
Grossnickle 2005b; Chirino et al. 2008; Cuesta et al. 2010a,b). Greater root growth in
larger seedlings after planting can reduce plant water stress and increase survival
(Hines and Long 1986; Luis et al. 2009).
Root system fibrosity
Greater root system fibrosity (i.e. branchiness) in bareroot seedlings (i.e. through under-
cutting—Faulkner 1953; Tanaka et al. 1976; Stupendick and Shepherd 1980; Schultz and
Thompson 1996) has been related to increased root growth capability (Rook 1969; Bacon
and Bachelard 1978; Hallgren and Tauer 1989) and in certain instances increased survival
(Shoulders 1959; Tanaka et al. 1976; Kormanik 1986; Hallgren and Tauer 1989; Muse and
Hatchell 1992; Schultz and Thompson 1996; Li et al. 2011). Greater root system fibrosity
can lead to greater water movement capability through the root system (Carlson 1986)
thereby reducing seedling water stress (Nambiar 1984). Conversely, stripping lateral roots
during the lifting of bareroot seedlings, thus reducing root system fibrosity and root growth
capability (South and Stumpff 1990), can reduce survival after field planting (Dierauf et al.
1992).
Manipulation of the container surface can increase root system fibrosity of container-
grown seedlings (i.e. copper treatment on container walls; Burdett and Martin 1982;
McDonald et al. 1984 and side slit containers for air pruning; Whitcomb 1984). These
container treatments redistribute primary roots more evenly along the plug length (Wenny
et al. 1988; Jones et al. 2002; Sword Sayer et al. 2011) and increase the number of first
order lateral roots (McDonald et al. 1982; Smith and Mccubbin 1992; Nelson 1999). This
can result in greater root growth capability (Arnold and Struve 1989; Dumroese 2000;
Moore 2002; Tsakaldimi and Ganatsas 2006) and increased seedling survival (Barnett and
McGilvray 1974).
Stocktype influence
Stocktype selection can influence survival because bareroot and container-grown seedlings
can have differing root growth capabilities. Extensive comparisons between container and
bareroot stocktypes across a range of sites lack no clear consensus favouring survivability
of one particular stocktype (Hobbs 1984; Owston 1990; Grossnickle 2000). Trials that
detected any survival differences, found container-grown, compared to bareroot, seedlings
had greater initial survival on droughty sites (Arnott 1981; Dixon et al. 1981; Hahn and
Smith 1983; Hobbs and Wearstler 1983; Burdett et al. 1984; South and Barnett 1986;
Becker et al. 1987; Barnett and McGilvray 1993; Nilsson and O
¨rlander 1995; Wilson et al.
2007).
Container-grown, compared to bareroot, seedlings can have greater root growth during
the first field growing season (Dixon et al. 1981; Burdett et al. 1984; Johnson et al. 1984;
Becker et al. 1987; Wilson et al. 2007). This improved root growth for container-grown
seedlings reduces their resistance to water flow through the soil–plant-atmosphere-con-
tinuum (Dixon et al. 1983; Grossnickle and Blake 1987) thereby minimizing plant water
stress (Burdett et al. 1984; Becker et al. 1987; Blake and Sutton 1987). Increased survival
724 New Forests (2012) 43:711–738
123
of container-grown, compared to bareroot, seedlings on drought prone sites has been
related to lower plant water stress (Hobbs and Wearstler 1983; Nilsson and O
¨rlander 1995).
Superior survival capability of container-grown, over bareroot, seedlings has also been
attributed to their intact, undisturbed and multidimensional root systems that minimizes
stress and leads to a quick root growth response (Tinus 1974). A growing media based plug
surrounding the root system can improve seedling water status, compared to bareroot
seedlings, independent from soil water availability (Jutras et al. 2007). Encasing roots of
container-grown seedlings in a plug acts as a protective barrier against root desiccation,
thereby creating a favorable plant water status to support photosynthesis and root growth
(Burdett 1990; Mena-Petite et al. 2001).
Restoration site performance
Root growth, though critical for survival and (or) growth of seedlings is not a perfect
predictor of seedling performance on forest restoration sites (Simpson and Ritchie 1997).
The relationship between root growth capability and field performance varies. Many
studies have shown that when seedlings grow roots they survive in the field (Stone 1955;
Sutton 1980; Burdett et al. 1983,1984; Feret and Kreh 1985; Hines and Long 1986; Larsen
et al. 1988; Simpson 1990; Simpson and Vyse 1995; McTague and Tinus 1996; Mena-
Petite et al. 2001; del Campo et al. 2007). Reviews on this issue (Ritchie and Dunlap
1980—26 trials, Ritchie and Tanaka 1990—12 trials) found *80 % of trials reported a
positive relationship between root growth capability and seedling survival. Though most
work has found a positive relationship between root growth and survival, the lack of
consistent trend has led to operational assessments questioning the importance of root
growth to seedling survival (South and Hallgren 1997). An extensive assessment of
operational plantings in British Columbia Canada, found that root growth potential failed
to display a strong relationship with field survival (Binder et al. 1988).
A reassessment of the Binder et al. (1988) data base using boundary line analysis might
reveal how root growth relates to seedling survival capability across a range of forest
restoration sites. Field data collected across a range of environmental conditions tends to
have a scatter of biological response to these conditions. On careful examination, normally
a series of biological response points occur within an upper boundary of response to the
dependent variable. Jarvis (1976) stated that this upper boundary is a maximum biological
response one can expect in the way of plant performance, while Webb (1972) credited the
scatter of biological response points below the boundary line to errors in measurement,
variability of biological data and the interaction with other site environmental factors.
Several researchers (Chambers et al. 1985; Grossnickle and Arnott 1992) used this premise
to create upper boundary biological response data sets (i.e. systematic selection of
20–25 % of data from across the dependent variable range) to quantify the maximum
biological response expected with a given dependent variable. When used on the Binder
et al. (1988) data, the boundary line shows that low root growth capability equates with the
chance of low survival, while high root growth capability equates with the chance of high
survival when these seedlings were field planted (Fig. 8). A further examination of this
data showed that if seedlings had low root growth capability (i.e. Index of Root Growth or
IRG \1) the probability was 52 % that the field plantation had \75 % survival, while
some seedling populations with low root growth capability had high field survival rates.
As the root growth capability of sample seedlings increased there was a greater chance of
survival. Seedlings with very high root growth capability (i.e. IRG [4) had \10 % chance
of a plantation having \75 % survival.
New Forests (2012) 43:711–738 725
123
Whether or not newly planted seedlings initially require new root growth for proper
field performance is related to the planting stress phenomenon. One way planting stress is
relieved is when root growth occurs and seedling water stress is reduced (Grossnickle
2005a). Simpson and Ritchie (1997) believe that root growth is strongly related to field
performance when seedlings have an inherently low level of stress resistance and/or when
site environmental conditions become more severe. These are conditions that lead to
planting stress. However, if seedlings are not exposed to planting stress, then initial root
growth is not essential for good field performance (Simpson and Ritchie 1997). This view
is exemplified by Stone et al. (2003), where critical root growth capacity (i.e. the minimum
root growth required for seedling survival on a given planting site) was twice as high for
harsh sites compared to gentle sites. Thus, survival of newly planted seedlings improves
when they can extend new roots into the soil, ensuring that water intake equals or exceeds
water loss (Wakeley 1948,1954) and forest restoration site conditions dictate the amount
of root growth required to overcome planting stress and ensure survival.
Conclusions
Morphological plant attributes influence a seedlings ability to survive after being planted
on to a forest restoration site because a plants susceptibility to drought induced mortality
can be due to their hydraulic architecture (McDowell et al. 2008). Thus, nursery cultural
practices that alter seedling morphological characteristics (i.e. hydraulic architecture) can
limit their susceptibility to planting stress (i.e. water stress). This is why morphological
attributes such as greater seedling stem diameter and root system size confer a higher
chance of survival. Seedling balance between the shoot and root systems, and seedling
overall size need to be adjusted in relation to potential forest restoration site environmental
conditions. Greater shoot system height is important if competition for light within the
Fig. 8 Second year seedling survival of field planted seedlings to mean index of root growth potential
(adapted from Binder et al. 1988). Data is from operational testing of 540 samples of seedlings. Data
includes 12 species and numerous stocktypes tested by the British Columbia Ministry of Forests. Mean
index root growth potential (IRG) classes are: 0no new root growth, 1some new roots, but none over 2 cm,
21–3 new roots over 1 cm long, 34–10 new roots over 1 cm long, 411–30 new roots over 1 cm long, 5and
6[35 new roots over 1 cm long. Boundary layer analysis (data defined by solid triangles) and the
regression equation were determined by analysis procedures of Grossnickle and Arnott (1992)
726 New Forests (2012) 43:711–738
123
vegetation complex is the potential site limiting factor. A somewhat smaller shoot system
and/or lower S:R are critical attributes if dry soils and high evaporative demand are
potential site limiting factors. However, morphological parameters are only measures that
help define overall seedling size, growth potential and balance, while seedling physio-
logical quality and root growth capability also have a major influence on survival.
Factors affecting seedling physiological quality such as stress resistance and nutritional
status have a major influence on survival. Improved survival is attributed to greater stress
resistance (i.e. through nursery cultural hardening practices that enhance drought resistance
and freezing tolerance) and improved seedling nutrition at planting (i.e. through fall nutrient
loading) that increases the speed with which seedlings can overcome planting stress and
become established on the forest restoration site. However, direct benefits from improved
seedling physiological attributes are ephemeral. Thus the primary purpose of these improved
physiological attributes is in aiding root growth, displaying a shoot system of sufficient size
within the forest restoration site complex and conferring improved seedling establishment
within months of planting. Ultimately the combination of a well-established seedling having
desirable shoot and root development is what ensures high survival.
Improved survival can be attributed to greater root growth immediately after seedlings
are field planted. Greater root growth is the result of greater root system size, fibrosity, plus
greater seedling stress resistance and improved nutrient status. Stocktype selection can also
influence survival, especially on droughty sites, because container-grown, compared to
bareroot, seedlings can have better root growth that is related to plug protection of roots
against desiccation and greater root growth capability that reduces potential planting stress.
These conclusions reiterate long held beliefs within the forest restoration community
that desirable morphological attributes (Toumey 1916), root growth (Stone 1955) and
physiological attributes (Wakeley 1948,1954; Tinus 1974) improve chances for increased
seedling survival. Work conducted during the past half century has confirmed that these
historical beliefs were correct, while defining species specific ranges for these plant
attributes. Planting seedlings with these desirable plant attributes does not guarantee high
survival; rather planting seedlings with desirable attributes increases chances for survival
within a forest restoration program.
Acknowledgments I thank Dr. Steve Colombo for the wide ranging discussions that lead to the initial
structure and focus of this paper. I also thank Drs. John Mexal and Dave South, and the Editor-in-Chief and
three anonymous reviewers whose review of draft versions of this manuscript kept me on topic and within
context of the published literature.
References
Abrams MD (1988) Sources of variation in osmotic potentials with special reference to North American tree
species. For Sci 34:1030–1046
Andivia E, Fernandez M, Va
´zquez-Pique
´J (2011) Autumn fertilization of Quercus ilex ssp. ballota (Desf.)
Samp. nursery seedlings: effects on morpho-physiology and field performance. Ann For Sci
68:543–553
Andivia E, Ma
´rquez-Garcı
´aB,Va
´zquez-Pique
´J, Co
´rdoba F, Fernandez M (2012) Autumn fertilization with
nitrogen improves nutritional status, cold hardiness and oxidative stress response of Holm oak
(Quercus ilex ssp. ballota (Desf.) Samp.) nursery seedlings. Trees 26:311–320
Armson KA, Sadreika V (1979) Forest nursery soil management and related practices. Ontario Ministry of
Natural Resources, Toronto, ON
Arnold MA, Struve DK (1989) Growing green ash and red oak in CuCO
3
-treated containers increases root
regeneration and shoot growth following transplant. J Am Soc Hort Sci 114:402–406
New Forests (2012) 43:711–738 727
123
Arnott JT (1981) Survival and growth of bullet, styroplug and bareroot seedlings on mid-elevation sites in
coastal British Columbia. For Chron 57:65–70
Arnott JT, Grossnickle SC, Puttonen P, Mitchell AK, Folk RS (1994) Influence of nursery culture on growth,
cold hardiness and drought resistance of yellow cypress. Can J For Res 23:2537–2547
Bacon GJ, Bachelard EP (1978) The influence of nursery conditioning treatments on some physiological
responses of recently transplanted seedlings of Pinus caribaea Mor. var. honduriensis B. and G. Aust
For Res 8:171–183
Baer N, Ronco F, Barney CW, Baer NW (1977) Effects of watering, shading and size of stock on survival of
planted lodgepole pine. USDA Forest Service Res Note RM-347
Baldwin VC, Barney CW (1976) Leaf water potential in planted ponderosa and lodgepole pines. For Sci
22:344–350
Barnett JP, McGilvray JM (1974) Copper screen controls root growth and increases survival on contain-
erized southern pine seedlings. Tree Planters’ Notes 25:11–12
Barnett JP, McGilvray JM (1993) Performance of container and bareroot loblolly pine seedlings on bot-
tomlands in South Carolina. South J Appl For 17:80–83
Bayley AD, Kietzka JW (1997) Stock quality and field performance of Pinus patula seedlings produced
under two nursery growing regimes during seven different nursery production periods. New For
13:341–356
Becker CA, Mroz GD, Fuller LG (1987) The effects of plant moisture stress on red pine (Pinus resinosa)
seedling growth and establishment. Can J For Res 17:813–820
Beikircher B, Florineth F, Mayr S (2010) Restoration of rocky slopes based on planted gabions and use of
drought-preconditioned woody specis. Ecol Eng 36:421–426
Benzian B, Brown RM, Freeman SCR (1974) Effect of late-season topdressing of N (and K) applied to conifer
transplants in the nursery on their survival and growth on British forest sites. Forestry 47:153–184
Bergquist J, O
¨rlander G (1998) Browsing damage by roe deer on Norway spruce seedlings planted on
clearcuts of different ages: 2. Effect of seedling vigour. For Ecol Manage 105:295–302
Bernier PY (1992) Soil texture influences seedling water stress in more ways than one. Tree Planters’ Notes
43:39–42
Bernier PY, Lamhamedi MS, Simpson DG (1995a) Shoot:root ratio is of limited use in evaluating the
quality of container conifer stock. Tree Planters’ Notes 46:102–106
Bernier PY, Stewart JD, Gonzalez A (1995b) Effects of the physical properties of sphagnum peat on water
stress in containerized Picea mariana seedlings under simulated field conditions. Scand J For Res
10:184–189
Biel C, Save
´R, Habrouk A, Espelta JM, Retana J (2004) Effects of restricted watering and CO
2
enrichment
in the morphology and performance after transplanting of nursery-grown Pinus nigra seedlings.
HortScience 39:535–540
Bigras FJ, Ryyppo A, Linderstrom A, Stattin E (2001) Cold acclimation and deacclimation of shoots and
roots of conifer seedlings. In: Bigras FJ, Colombo SJ (eds) Conifer cold hardiness. Kluwer, The
Netherlands, pp 57–88
Binder WD, Scagel RK, Krumlik GJ (1988) Root growth potential: facts, myths, value? USDA Forest
Service Gen. Tech. Rep. RM-167, pp 111–118
Binkley D (1986) Forest nutrition management. Wiley, New York
Birchler TM, Rose R, Haase DL (2001) Fall fertilization with N and K: effects on Douglas-fir seedlings
quality and performance. West J Appl For 16:71–79
Blake TJ, Sutton RF (1987) Variation in water relations of black spruce stock types planted in Ontario. Tree
Physiol 3:331–344
Blake J, Zaerr J, Hee S (1979) Controlled moisture stress to improve cold hardiness and morphology of
Douglas-fir seedlings. For Sci 25:576–582
Blake JL, Teeter LD, South DB (1989) Analysis of economic benefits from increasing uniformity in
Douglas-fir nursery stock. In: Mason WL, Deans JD, Thompson S (eds) Producing uniform conifer
planting stock, vol 26, pp 251–262 (for suppl)
Boivin JR, Salifu KF, Timmer VR (2004) Late season fertilization of Picea mariana seedlings: intensive
loading and outplanting response on greenhouse bioassays. Ann For Sci 61:737–745
Boyer JN, South DB (1987) Excessive seedling height, high shoot-to-root ratio and benomyl dip reduce
survival of stored loblolly pine seedlings. Tree Planters’ Notes 37:19–21
Brissette JC, Roberts TC (1984) Seedling size and lifting date effects on root growth potential of loblolly
pine from two Arkansas nurseries. Tree Planters’ Notes 35:34–38
Brix H, van den Driessche R (1974) Mineral nutrition of container-grown tree seedlings. In: Tinus RW,
Stein WI, Balmer WE (eds) Proceedings of the North American containerized forest tree seedling
symposium. Great Plains Ag. Council Publ. No. 68, pp 77–84
728 New Forests (2012) 43:711–738
123
Burdett AN (1987) Understanding root growth capacity: theoretical considerations in assessing planting
stock quality by means of root growth tests. Can J For Res 17:768–775
Burdett AN (1990) Physiological processes in plantation establishment and the development of specifica-
tions for forest planting stock. Can J For Res 20:415–427
Burdett AN, Martin PAF (1982) Chemical root pruning of coniferous seedlings. HortScience 17:622–624
Burdett AN, Simpson DG, Thompson CF (1983) Root development and plantation establishment success.
Plant Soil 71:103–110
Burdett AN, Herring LJ, Thompson CF (1984) Early growth of planted spruce. Can J For Res 14:644–651
Burney OT, Jacobs DF (2011) Ungulate herbivory of regenerating conifers in relation to foliar nutrition and
terpenoid production. For Ecol Manage 262:1834–1845
Burr KE (1990) The target seedling concept: bud dormancy and cold-hardiness. In: Rose R, Campbell SJ,
Landis TD (eds) Target seedling symposium: proceedings of the western forest nursery associations.
USDA Forest Service Gen. Tech. Rep. RM-200, pp 79–90
Buxton GF, Cyr DR, Dumbroff EB (1985) Physiological responses of three northern conifers to rapid and
slow induction of moisture stress. Can J Bot 63:1171–1176
Calme
´S, Margolis HA, Bigras FJ (1993) Influence of cultural practices on the relationship between frost
tolerance and water content of containerized black spruce, white spruce, and jack pine seedlings. Can J
For Res 23:503–511
Carlson WC (1986) Root system considerations in the quality of loblolly pine seedlings. South J Appl For
10:87–92
Chambers JL, Hinckley TM, Cox GS, Metcalf CL, Aslin RG (1985) Boundary-line analysis and models of
leaf conductance for four oak-hickory forest species. For Sci 31:437–450
Chamshama SAO, Hall JB (1987) Effects of nursery treatments on Eucalyptus camaldulensis field estab-
lishment and early growth at Mafiga, Morogoro, Tanzania. For Ecol Manage 21:91–108
Chirino E, Vilagrosa A, Herna
´ndez EI, Matoc A, Vallejoa VR (2008) Effects of deep container on morpho-
fuctional characteristics and root colonization in Quercus suber L. seedlings for reforestation in
Mediterranean climate. For Ecol Manage 256:779–785
Christersson L (1972) The transpiration rate of unhardened, hardened, and dehardened seedlings of spruce
and pine. Physiol Plant 26:258–263
Cleary BD, Greaves RD, Hermann RK (eds) (1978a) Regenerating Oregon’s forests: a guide for the
regeneration forester. Oregon State University Extension Service, Corvallis, OR
Cleary BD, Greaves RD, Owsten PW (1978b) Seedlings. In: Cleary BD, Greaves RD, Hermann RK (eds)
Regenerating Oregon’s forests: a guide for the regeneration forester. Oregon State University Exten-
sion Service, Corvallis, OR, pp 63–97
Clemens J, Jones PG (1978) Modification of drought resistance by water stress conditioning in Acacia and
Eucalyptus. J Exp Bot 29:895–904
Close DC, McArthur C, Pietrzykowski E, Fitzgerald H, Paterson S (2004) Evaluating effects of nursery and
post-planting nutrient regimes on leaf chemistry and browsing of eucalypt seedlings in plantations. For
Ecol Manage 200:101–112
Close DC, Bail I, Hunter S, Beadle CL (2005) Effects of exponential nutrient-loading on morphological and
nitrogen characteristics and on after planting performance of Eucalyptus globulus seedlings. For Ecol
Manage 205:397–403
Colombo SJ (1987) Changes in osmotic potential, cell elasticity, and turgor relationships of 2nd-year black
spruce container seedlings. Can J For Res 17:365–369
Colombo SJ, Menzies MI, O’Reilly C (2001) Influence of nursery cultural practices on cold hardiness of
coniferous forest tree seedlings. In: Bigras FJ, Colombo SJ (eds) Conifer cold hardiness. Kluwer, The
Netherlands, pp 223–252
Colombo SJ, Glerum C, Webb DP (2003) Daylength, temperature and fertilization effects on desiccation
resistance, cold hardiness and root growth potential of Picea mariana seedlings. Ann For Sci
60:307–317
Cuesta B, Vega J, Villar-Salvador P, Rey-Benayas JM (2010a) Root growth dynamics of allepo pine (Pinus
halipensis Mill.) seedlings in relation to shoot elongation, plant size and tissue nitrogen concentration.
Trees 24:899–908
Cuesta B, Vega J, Villar-Salvador P, Pue
´rtolas J, Jacobs DF, Rey-Benayas JM (2010b) Why do large,
nitrogen rich seedlings better resist stressful transplanting conditions? A physiological analysis in two
contrasting Mediterranean forests. For Ecol Manage 20:71–78
D’Aoust AL, Cameron SE (1982) The effects of dormancy induction, low temperature and moisture stress
on cold hardening of containerized black spruce seedlings. In: Scarrett JB, Glerum C, Plexman CA
(eds) Proceedings of the Canadian containerized tree seedling symposium. Can. For. Serv. Great Lakes
For. Cent. COJFRC Symp. Proc. O-P-10, pp 153–161
New Forests (2012) 43:711–738 729
123
Davis AD, Jacobs DF (2005) Quantifying root system quality of nursery seedlings and relationship to
outplanting performance. New For 30:295–311
Day RJ, Skoupy J (1971) Moisture storage capacity and post-planting patterns of moisture movement from
seedling containers. Can J For Res 1:151–158
del Campo AD, Navarro RM, Hermoso J, Iba
´n
´ez AJ (2007) Relationship between root growth potential and
field performance in Aleppo pine. Ann For Sci 64:541–548
del Campo AD, Navarro RM, Ceacero CJ (2010) Seedling quality and field performance of commercial
stocklots of containerized holm oak (Quercus ilex) in Mediterranean Spain: an approach for estab-
lishing a quality standard. New For 39:19–37
Dierauf TD, Chandler LA, Hixson DL (1992) Stripping of lateral roots from loblolly pine seedlings during
lifting—effect on field performance. Virginia Dept. For. Occ. Rep. 105
Dixon RK, Garrett HE, Cox GS, Johnson PS, Sander IL (1981) Container- and nursery-grown black oak
seedlings inoculated with Pisolithus tinctorius: growth and ectomycorrhizal development following
outplanting on an Ozark clear-cut. Can J For Res 11:492–496
Dixon RK, Pallardy SG, Garrett HE, Cox GS (1983) Comparative water relations of container-grown and
bare-root ectomycorrhizal and nonmycorrhizal Quercus velutina seedlings. Can J Bot 61:1559–1565
Dumroese KR (2000) Changes in interior Douglas-fir root development in containers after copper and auxin
treatments. West J Appl For 15:213–216
Duryea ML (1984) Nursery cultural practices: impacts on seedling quality. In: Duryea ML, Landis TD (eds)
Forest nursery manual: production of bareroot seedlings. Martinus Nijhoff/Dr. W. Junk Publishers, The
Hague, pp 143–164
Duryea ML, Dougherty PM (eds) (1991) Forest regeneration manual. Kluwer, Dordrecht
Duryea ML, McClain KM (1984) Altering seedling physiology to improve reforestation success. Corvallis,
OR. In: Duryea ML, Brown GH (eds) Seedling physiology and reforestation success: proceedings of the
physiology working group technicalsession. Martinus Nijhoff/Dr. W. Junk Publishers, Boston, pp 77–114
Faulkner R (1953) Early observations on the root development of one-year-old Corsican pine seedlings
following root pruning. Scott For 7:23–26
Feret RP, Kreh RE (1985) Seedling root growth potential as an indicator of loblolly pine field performance.
For Sci 31:1005–1011
Floistad IS, Kohmann K (2004) Influence of nutrient supply spring frost hardiness and time of bud break in
Norway spruce (Picea abies (L.) Karst.) seedlings. New For 27:1–11
Foiles MW, Curtis JD (1973) Regeneration of ponderosa pine in the northern Rocky Mountain-Inter-
mountain region. USDA Forest Service Res. Paper INT-145, 45 pp
Folk RS, Grossnickle SC, Major JE, Arnott JT (1994) Influence of nursery culture on western red cedar. II.
Freezing tolerance of fall-planted seedlings and morphological development of fall- and spring-planted
seedlings. New For 8:231–247
Fuchigami LH, Nee CC (1987) Degree growth stage model and rest-breaking mechanisms in temperate
woody perennials. HortScience 22:836–845
Ge
´ne
´re
´B, Garriou D (1999) Stock quality and field performance of Douglas-fir seedlings under varying
degrees of water stress. Ann For Sci 56:501–510
Gleason JF, Duryea M, Rose R, Atkinson M (1990) Nursery and field fertilization of 2?0 ponderosa pine
seedlings: the effect on morphology, physiology and field performance. Can J For Res 20:1766–1772
Grossnickle SC (2000) Ecophysiology of northern spruce species: the performance of planted seedlings.
NRC Research Press, Ottawa
Grossnickle SC (2005a) Importance of root growth in overcoming planting stress. New For 30:273–294
Grossnickle SC (2005b) Seedling size and reforestation success. How big is big enough? In: Colombo SJ
(Compiler) The thin green line: a symposium on the state-of-the-art in reforestation. Ont. For. Res.
Inst., Ontario Ministry of Natural Resources. For. Res. Info. Paper 160, pp 138–144
Grossnickle SC, Arnott JT (1992) Gas exchange response of western hemlock seedlings from various
dormancy induction treatments to reforestation site environmental conditions. For Ecol Manage
49:177–193
Grossnickle SC, Blake TJ (1987) Comparison of water relation patterns for newly planted bare-root and
container jack pine and black spruce seedlings on boreal cut-over sites. New For 1:101–116
Grossnickle SC, Folk RS (2003) Spring versus summer spruce stocktypes of western Canada: nursery
development and field performance. West J Appl For 18:267–275
Grossnickle SC, Folk RS (2007) Field performance potential of a somatic interior spruce seedlot. New For
34:51–72
Grossnickle SC, Major JE (1994) Interior spruce seedlings compared to emblings produced from somatic
embryogenesis. III. Physiological response and morphological development on a reforestation site. Can
J For Res 24:1397–1407
730 New Forests (2012) 43:711–738
123
Grossnickle SC, Reid CPP (1984) Water relations of Engelmann spruce seedlings on a high-elevation mine
site: an example of how reclamation techniques can alter microclimate and edaphic conditions. Reclam
Reveg Res 3:199–221
Grossnickle SC, Arnott JT, Major JE, Tschaplinski TJ (1991a) Influence of dormancy induction treatment on
western hemlock seedlings. 1) Seedling development and stock quality assessment. Can J For Res
21:164–174
Grossnickle SC, Major JE, Arnott JT, LeMay VM (1991b) Stock quality assessment through an integrated
approach. New For 5:77–91
Grossnickle SC, Arnott JT, Major JE (1991c) Influence of dormancy induction treatments on western
hemlock seedlings. 2) Physiological and morphological response during the first growing season on a
reforestation site. Can J For Res 21:175–185
Guehl JM, Aussenac G, Kaushal P (1989) The effects of transplanting stress on photosynthesis, stomatal
conductance and leaf water potential in Cedrus atlantica: role of root regeneration. Ann Sci For
46S:464–468
Haase DL, Rose R (1993) Soil moisture stress induces transplant shock in stored and unstored 2?0 Douglas-
fir seedlings of varying root volumes. For Sci 39:275–294
Hahn PF, Smith AJ (1983) Douglas-fir planting stock performance. Comparison after the third growing
season. Tree Planters’ Notes 34:33–39
Hallgren SW, Tauer CG (1989) Root growth potential, first-year survival, and growth of shortleaf pine
seedlings show effects of lift date, storage, and family. South J Appl For 13:163–169
Harvey HP, van den Driessche R (1997) Nutrition, xylem cavitation and drought resistance in hybrid
poplars. Tree Physiol 17:647–654
Harvey HP, van den Driessche R (1999) Nitrogen and potassium on xylem cavitation and water-use
efficiency in poplars. Tree Physiol 19:943–950
Hennessey TC, Dougherty PM (1984) Characterization of the internal water relations of loblolly pine
seedlings in response to nursery cultural treatments: implications for forest regeneration success. In:
Duryea ML, Brown GN (eds) Seedling physiology and reforestation success. Martinus Nijhoff Dr W.
Junk Publishers, Dordrecht, pp 225–244
Hermann RK (1964) Importance of top-root ratios for survival of Douglas-fir seedlings. Tree Planters’ Notes
64:7–11
Hines FD, Long JN (1986) First-and second-year survival of containerized Engelmann spruce in relation to
initial seedling size. Can J For Res 16:668–670
Hobbs SD (1982) Stocktype selection and planting techniques for Douglas-fir on skeletal soils in southwest
Oregon. In: Hobbs SD, Helgerson OT (eds) Proceeding of reforestation of skelatal soils workshop.
Forest Research Laboratory, Oregon State University, Corvallis, pp 92–96
Hobbs SD (1984) The influence of species and stocktype selection on stand establishment: an ecophysio-
logical perspective. In: Duryea ML, Brown GH (eds) Seedling physiology and reforestation success:
proceedings of the physiology working group technical session. Martinus Nijhoff/Dr. W. Junk Pub-
lishers, Boston, pp 180–224
Hobbs SD, Wearstler KA (1983) Performance of three Douglas-fir stocktypes on a skeletal soil. Tree
Planters’ Notes 34:11–14
Hobbs SD, Crawford MS, Yelczyn BA (1989) Early development of three Douglas-fir stocktypes on a
droughty skeletal soil. West J Appl For 4:21–24
Hobbs SD, Tesch SD, Owston PW, Stewart RE, Tappeiner JC II, Wells GE (1992) Reforestation practices in
Southwestern Oregon and Northern California. Forest Research Laboratory, Oregon State University,
Corvallis, OR
Ingestad T, Lund AB (1986) Theory and techniques for steady state mineral nutrition and growth of plants.
Scand J For Res 1:439–453
Irwin KM, Duryea ML, Stone EL (1998) Fall-applied nitrogen improves performance of 1-0 slash pine
nursery seedlings after outplanting. South J Appl For 22:111–116
Islam MA, Apostol KG, Jacobs DF, Dumroese RK (2009) Fall fertilization of Pinus resinosa seedlings:
nutrient uptake, cold hardiness, and morphological development. Ann For Sci 66:704–713
Jacobs DF, Davis AD, Wilson BC, Dumroese RK, Goodman RC, Salifu KF (2008) Short-day treatment
alters Douglas-fir seedling dehardening and transplant root proliferation at varying rhizosphere tem-
peratures. Can J For Res 38:1526–1535
Jarvis PG (1976) The interpretation of the variations in leaf water potential and stomatal conductance found
in canopies in the field. Philos Trans R Soc Lond Ser B 273:593–610
Jobidon R, Charette L, Bernier PY (1997) Initial size and competing vegetation effects on water stress and
growth of Picea mariana (Mill.) BSP seedlings planted in three different environments. For Ecol
Manage 103:295–308
New Forests (2012) 43:711–738 731
123
Jobidon R, Roy V, Cyr G (2003) Net effect of competing vegetation on selected environmental conditions
and performance of four spruce seedling stock sizes after eight years in Quebec (Canada). Ann For Sci
60:691–699
Johnsen KH, Feret PP, Seiler JR (1988) Root growth potential and shoot activity of northern and southern
provenances of 1-0 eastern white pine seedlings grown in a Virginia nursery. Can J For Res
18:610–614
Johnson JD, Cline ML (1991) Seedling quality of southern pines. In: Duryea ML, Dougherty PM (eds)
Forest regeneration manual. Kluwer, Dordrecht, pp 143–162
Johnson DM, Smith WK (2005) Refugial forests of the southern Appalachians: photosynthesis and survival
in current-year Abies fraseri seedlings. Tree Physiol 25:1379–1387
Johnson PS, Novinger SL, Mares WG (1984) Root, shoot, and leaf area growth potentials of nortern red oak
planting stock. For Sci 30:1017–1026
Johnson JD, Seiler JR, McNabb KL (1985) Manipulation of pine seedling physiology by water stress
conditioning. In: South DB (ed) Proceedings of the international symposium on nursery manage.
Practices for the Southern Pines, IUFRO Sub. Grp. S3.202-03, pp 290–302
Jones MD, Kiiskila S, Flanagan A (2002) Field performance of pine stock types: two-year results of a trial
on interior lodgepole pine seedlings grown in Styroblocks
TM
, Copperblocks
TM
, or AirBlocks
TM
.BCJ
Ecosyst Manag 2:1–12
Jospon TM, Paul JL (1985) Influence of fall fertilization and moisture stress on growth and field performance
on container-grown Douglas-fir seedlings. USDA Forest Service Gen. Tech. Rep. INT-185, pp 14–19
Jutras S, Thiffault N, Munson AD (2007) Comparing large bareroot and container stock: water stress as
influenced by peat and soil water availability. Tree Planters’ Notes 52:15–18
Kainer KA, Duryea ML (1990) Root wrenching and lifting date of slash pine: effects on morphology,
survival and growth. New For 4:207–221
Kandiko RA, Timmis R, Worrall J (1980) Pressure-volume curves of shoots and roots of normal and
drought-conditioned western hemlock seedlings. Can J For Res 10:10–16
Kaushal P, Aussenec G (1989) Drought preconditioning of Corsican pine and Cedar of Atlas seedlings:
photosynthesis, transpiration and root regeneration after transplanting. Acta Oecol 11:61–78
Kim YT, Colombo SJ, Hickie DF, Noland TD (1999) Amino acid, carbohydrate, glutathione, mineral
nutrient and water potential changes in non-water-stressed Picea mariana seedlings after transplanting.
Scand J For Res 14:416–424
Kittredge J (1929) Forest planting in the lake states. USDA Bull. 1497
Kormanik PP (1986) Lateral root morphology as an expression of sweetgum seedling quality. For Sci
32:595–604
Kozlowski TT, Pallardy SG (2002) Acclimation and adaptive response of woody plants to environmental
stress. Bot Rev 68:270–334
Kozlowski TT, Kramer PJ, Pallardy SG (1991) The physiological ecology of woody plants. Academic Press,
New York
Krasowski MJ, Letchford T, Eastham AM (1993) Growth of short-day treated spruce seedlings planted
throughout British Columbia. Forestry Canada and British Columbia Ministry of Forests, Victoria, BC,
FRDA Rep. 209
Lamhamedi MS, Bernier PY, He
´rbert C (1997) Effect of shoot size on the gas exchange and growth of
containerized Picea mariana seedlings under different watering regimes. New For 13:209–223
Landis TD, Tinus RW, McDonald SE, Barnett JP (1989) Seedling nutrition and irrigation. The container tree
nursery manual, vol 4. USDA Forest Service Agric. Handb. 674, Washington, DC, 87 pp
Landis TD, Tinus RW, Barnett JP (1999) Seedling propagation. The container tree nursery manual, vol. 6.
USDA Forest Service Agric. Handb. 674, Washington, DC, 167 pp
Lantz CW (ed) (1985) Southern pine nursery handbook. USDA Forest Service Cooperative Forestry,
Southern Region
Larsen HS, South DB, Boyer JM (1986) Root growth potential, seedling morphology and bud dormancy
correlate with survival of loblolly pine seedlings planted in December in Alabama. Tree Physiol
1:253–263
Larsen HS, South DB, Boyer JM (1988) Foliar nitrogen content at lifting correlates with early growth of
loblolly pine seedlings from 20 nurseries. South J Appl For 12:181–185
Lavender DP (1985) Bud dormancy. In: Duryea ML (ed) Evaluating seedling quality: principles, procedures,
and predictive ability of major tests. Forest Research Laboratory, Oregon State University, Corvallis,
OR, pp 7–15
Lavender DP, Cleary BD (1974) Coniferous seedling production techniques to improve seedling estab-
lishment. In: Tinus RW, Stein WI, Balmer WE (eds) Proceedings of the North American containerized
forest tree seedling symposium. Great Plains Ag. Council Publ. No. 68, pp 177–180
732 New Forests (2012) 43:711–738
123
Lavender DP, Wareing PF (1972) The effects of daylength and chilling on the response of Douglas-fir
(Pseudotsuga menziesii (Mirb.) Franco.) seedlings to root damage and storage. New Phytol
71:1055–1067
Lavender DP, Parish R, Johnson CM, Montgomery G, Vyse A, Willis RA, Winston D (eds) (1990)
Regenerating British Columbia’s forests. University of British Columbia Press, Vancouver, BC
Li GL, Lui Y, Yang J, Sun HY, Jia ZK, Ma LY (2011) Influence of initial age and size on the field
performance of Larix olgensis seedlings. New For 42:215–226
Long AJ, Carrier BD (1993) Effects of Douglas-fir 2?0 seedling morphology on field performance. New For
7:19–32
Lopushinsky W, Beebe T (1976) Relationship of shoot-root ratio to survival and growth of outplanted
Douglas-fir and ponderosa pine seedlings. USDA Forest Service Res. Note, PNW-274
Luis VC, Pue
´rtolas J, Climent J, Peters J, Gonza
´lez-Rodrı
´guez AM, Morales D, Jimene
´z MS (2009) Nursery
fertilization enhances survival and physiological status in Canary Island pine (Pinus canariensis)
seedlings planted in a semiarid environment. Eur J For Res 128:221–229
Luoranen J, Lahti M, Rikala R (2008) Frost hardiness of nutrient-loaded two-year-old Picea abies seedlings
in autumn and at the end of freezer storage. New For 35:207–220
Macey DE, Arnott JT (1986) The effect of moderate moisture and nutrient stress on bud formation and
growth of container-grown white spruce seedlings. Can J For Res 16:949–954
Major JE, Grossnickle SC, Folk RS, Arnott JT (1994) Influence of nursery culture on western red cedar.
I. Measurement of seedling attributes before fall and spring planting. New For 8:211–229
Malik V, Timmer VR (1996) Growth, nutrient dynamics, and interspecific competition of nutrient-loaded
black spruce seedlings on a boreal mixedwood site. Can J For Res 26:1651–1659
Malik V, Timmer VR (1998) Biomass partitioning and nitrogen retranslocation in black spruce seedlings on
competitive mixedwood sites: a bioassay study. Can J For Res 28:206–215
Margolis HA, Brand DG (1990) An ecophysiological basis for understanding plantation establishment. Can
J For Res 20:375–390
Margolis HA, Waring RH (1986) Carbon and nitrogen allocation patterns of Douglas-fir seedlings fertilized
with nitrogen in autumn. II. Field performance. Can J For Res 16:903–909
Marshall JD (1985) Carbohydrate status as a measure of seedling quality. In: Duryea ML (ed) Evaluating
seedling quality: principles, procedures, and predictive ability of major tests. Forest Research Labo-
ratory, Oregon State University, Corvallis, OR, pp 49–56
Mason EG, South DB, Weizhong Z (1996) Performance of Pinus radiata in relation to seedling grade, weed
control, and soil cultivation in the central North Island of New Zealand. N Z J For Sci 26:173–183
Mattsson A (1996) Predicting field performance using seedling quality assessment. New For 13:223–248
McDonald SE, Tinus RW, Reid CPP (1982) Root development control measures in containers: recent
findings. In: Scarrett JB, Glerum C, Plexman CA (eds) Proceedings of the Canadian containerized tree
seedling symposium. Can. For. Serv. Great Lakes For. Cent. COJFRC Symp. Proc. O-P-10,
pp 207–214
McDonald SE, Tinus RW, Reid CPP, Grossnickle SC (1984) Effect of CuCO
3
container wall treatments and
mycorrhizae fungi inoculation of growing medium on pine seedling growth and root development.
J Environ Hortic 2:5–8
McDowell N, Pockman WT, Allen CD, Bershears DD, Cobb N, Kolb T, Plaut J, Sperry J, West A, Williams
DG, Yepez EA (2008) Mechanisms of plant survival and mortality during drought: why do some plants
survive while others succumb to drought? New Phytol 178:719–739
McGrath DA, Duryea ML (1994) Initial moisture stress, budbreak and two-year field performance of three
morphological grades of slash pine seedlings. New For 8:335–350
McKay HM, Mason WL (1991) Physiological indicators of tolerance to cold storage in Sitka spruce and
Douglas-fir seedlings. Can J For Res 21:890–910
McTague JP, Tinus RW (1996) The effects of seedling quality and forest site weather on field survival of
ponderosa pine. Tree Planters’ Notes 47:16–32
Mena-Petite A, Ortega-Lasuen U, Gonza
´lez-Moro M, Lacuesta M, Mun
ˇoz-Rueda A (2001) Storage duration
and temperature effects on the functional integrity of container and bare-root Pinus radiata D. Don
stock-types. Trees 15:289–296
Menzies MI, Holden DG, Green LM, Rook DA (1981) Seasonal changes in frost tolerance of Pinus radiata
seedlings raised in different nurseries. N Z J For Sci 11:100–111
Menzies MI, van Dorsser JC, Balneaves JM (1985) Seedling quality—radiata pine as a case study. In: South
DB (ed) Proceedings of the international symposium on nursery manage. Practices for the Southern
Pines, IUFRO Sub. Grp. S3.202-03, pp 385–415
Mexal JG, Dougherty PM (1983) Growth of loblolly pine seedlings. IV. Performance in a simulated drought
environment. Weyerhaeuser Co. Hot Springs, AK. Tech Rep. 050-1422/6, 26 pp
New Forests (2012) 43:711–738 733
123
Mexal JG, Landis TD (1990) Target seedling concepts: height and diameter. In: Rose R, Campbell SJ,
Landis TD (eds) Target seedling symposium: proceedings of the western forest nursery associations.
USDA Forest Service Gen. Tech. Rep. RM-200, pp 17–36
Mexal JG, South DB (1991) Bareroot seedling culture. In: Duryea ML, Dougherty PM (eds) Forest
regeneration manual. Kluwer, Dordrecht, pp 89–115
Mexal JG, Timmis R, Morris WG (1979) Cold hardiness of containerized loblolly pine seedlings. South J
Appl For 3:15–19
Mexal JG, Cuevas Rangel RA, Landis TD (2008) Reforestation success in central Mexico: factors deter-
mining survival and early growth. Tree Planters’ Notes 53:16–22
Miller PC (1983) Comparison of water balance characteristics of plant species in ‘‘natural’’ versus modified
ecosystems. In: Mooney HA, Godron M (eds) Disturbance and ecosystems. Components of response.
Springer, Berlin, pp 188–212
Mohammed GH, Noland TL, Wagner RG (1998) Physiological perturbation in jack pine (Pinus banksiana
Lamb.) in the presence of competing herbaceous vegetation. For Ecol Manage 103:77–85
Moore DG (2002) Some new research into container design. Proc Int Plant Prop Soc 52:105–108
Morrissey RC, Jacobs DF, Davis AS, Rathfon RA (2010) Survival and competitiveness of Quercus rubra
regeneration associated with planting stocktype and harvest opening intensity. New For 40:273–287
Munson AD, Bernier PY (1993) Comparing natural and planted black spruce seedlings. II. Nutrient uptake
and efficiency of use. Can J For Res 23:2435–2442
Muse DH, Hatchell GE (1992) A preliminary identification of morphological indicators of field performance
in bare-root nursery stock. USDA Forest Service Res. Pap. SE-283
Nambiar EKS (1984) Significance of first-order lateral roots on the growth of young radiata pine under
environmental stress. Aust For Res 14:187–199
Nelson WR (1999) Root pruning can influence first order lateral root development of containerised plants.
Proc Int Plant Prop Soc 49:96–103
Newton M, Cole EC, White DE (1993) Tall planting stock for enhanced growth and domination of brush in
the Douglas-fir region. New For 7:107–121
Nilsson U, O
¨rlander G (1995) Effects of regeneration methods on drought damage to newly planted Norway
spruce seedlings. Can J For Res 25:790–802
Oliet JA, Tejada M, Salifu KF, Collazos A, Jacobs DF (2009a) Performance and nutrient dynamics of holm
oak (Quercus ilex L.) seedlings in relation to nursery nutrient loading and post-transplant fertility. Eur J
Forest Res 128:253–263
Oliet JA, Planelles R, Artero F, Valverde R, Jacobs DF, Segura M (2009b) Field performance of Pinus
halepensis planted in Mediterranean arid conditions: relative influence of seedling morphology and
mineral nutrition. New For 37:313–331
Oliet JA, Salazar JM, Villar R, Robredo E, Valladares F (2011) Fall fertilization of Holm oak affects N and
P dynamics, root growth potential, and post-planting phenology and growth. Ann For Sci 68:647–656
O’Reilly C, Owens JN, Arnott JT, Dunsworth BG (1994) Effect of nursery culture on morphological
development of western hemlock seedlings during field establishment. II. Survival, shoot length
components, and needle length. Can J For Res 24:61–70
O
¨rlander G, Due K (1986) Location of hydraulic resistance in the soil-plant pathway in seedlings of Pinus
sylvestris L. grown in peat. Can J For Res 16:115–123
Owston PW (1990) Target seedling specifications: Are stocktype designations useful? In: Rose R, Campbell
SJ, Landis TD (eds) Target seedling symposium: proceedings of the western forest nursery associa-
tions. USDA Forest Service Gen. Tech. Rep. RM-200, pp 9–16
Parker J (1949) Effects of variation in the root-leaf ratio on transpiration rate. Plant Physiol 24:739–743
Paul GS, Montagnini F, Berlyn GP, Craven DJ, van Breugel M, Hall JS (2012) Foliar herbivory and leaf
traits of five native tree species in a young plantation of Central Panama. New For 43:9–87
Pue
´rtolas J, Gil L, Pardos JA (2003) Effects of nutritional status and seedling size on field performance of
Pinus halepensis planted in former arable land in the Mediterranean basin. Forestry 76:159–168
Puttonen P (1997) Looking for the ‘‘silver bullet’’—can one test do it all? New For 13:9–27
Ritchie GA (1982) Carbohydrate reserves and root growth potential in Douglas-fir seedlings before and after
cold storage. Can J For Res 12:905–912
Ritchie GA (1984) Assessing seedling quality. In: Duryea ML, Landis TD (eds) Forest nursery manual:
production of bareroot seedlings. Martinus Nijhoff/Dr. W. Junk Publishers, The Hague, pp 243–266
Ritchie GA (1985) Root growth potential: principles, procedures, and predictive ability. In: Duryea ML (ed)
Evaluating seedling quality: principles, procedures, and predictive ability of major tests. Oregon State
University, Forestry Research Laboratory, Corvallis, OR, pp 93–106
Ritchie GA, Dunlap JR (1980) Root growth potential: its development and expression in forest tree seed-
lings. N Z J For Sci 10:218–248
734 New Forests (2012) 43:711–738
123
Ritchie GA, Roden JR (1985) Comparison between two methods of generating pressure-volume curves.
Plant, Cell Environ 8:49–53
Ritchie GA, Tanaka Y (1990) Root growth potential and the target seedling. In: Rose R, Campbell SJ,
Landis TD (eds) Target seedling symposium: proceedings of the western forest nursery associations.
USDA Forest Service Gen. Tech. Rep. RM-200, pp 37–51
Roberts DR, Dumbroff EB (1986) Drought resistance, transpiration rates and ABA levels of three northern
conifers. Tree Physiol 1:161–178
Rodgers AR, Williams D, Sinclair ARE, Sullivan TP, Andersen RJ (1993) Does nursery production reduce
antiherbivore defenses of white spruce? Evidence from feeding experiments with snowshoe hares. Can
J For Res 23:2358–2361
Rook DA (1969) Water relations of wrenched and unwrenched Pinus radiata seedlings on being trans-
planted into conditions of water stress. N Z J For 14:50–58
Rose R, Ketchum JS (2003) Interaction of initial seedling diameter, fertilization and weed control on
Douglas-fir growth over the first four years after planting. Ann For Sci 60:625–635
Rose R, Gleason JF, Atkinson M (1993) Morphological and water-stress characteristics of three Douglas-fir
stocktypes in relation to seedling performance under different soil moisture conditions. New For
7:1–17
Rose R, Haase DL, Kroiher F, Sabin T (1997) Root volume and growth of ponderosa pine and Douglas-fir
seedlings: a summary of eight growing seasons. West J Appl For 12:69–73
Royo A, Gil L, Pardos JA (2001) Effects of water stress conditioning on morphology, physiology and field
performance of Pinus halepensis Mill. seedlings. New For 21:127–140
Rowe SJ (1964) Environmental preconditioning with special reference to forestry. Ecology 45:399–403
Rudolf PO (1939) Why forest plantations fail. J For 37:377–383
Salifu KF, Jacobs DF, Birge ZKD (2009) Nursery nitrogen loading improves field performance of bareroot
oak seedlings planted on abandon mine lands. Restor Ecol 17:339–349
Scagel R, Bowden R, Madill M, Kooistra C (1993) Provincial seedling stock type selection and ordering
guidelines. British Columbia Ministry of Forests, Victoria, BC 75
Schultz RC, Thompson JR (1996) Effect of density control and undercutting on root morphology of 1?0
bareroot hardwood seedlings: five-year field performance of root-graded stock in central USA. New
For 13:297–310
Seiler JR, Johnson JD (1985) Photosynthesis and transpiration of loblolly pine seedlings as influenced by
moisture-stress conditioning. For Sci 31:742–749
Shoulders E (1959) Root pruning boosts longleaf pine survival. Tree Planters’ Notes 36:15–19
Simpson DG (1990) Frost hardiness, root growth capacity, and field performance relationships in interior
spruce, lodgepole pine, Douglas-fir, and western hemlock seedlings. Can J For Res 20:566–572
Simpson DG, Ritchie GA (1997) Does RGP predict field performance? A debate. New For 13:253–277
Simpson DG, Vyse A (1995) Planting stock performance: site and RGP effects. For Chron 71:739–742
Smith IE, McCubbin PD (1992) Effect of copper tray treatment on Eucalyptus grandis (Hill ex Maiden)
seedling growth. Acta Hortic 319:371–376
South DB (1993) Rationale for growing southern pine seedlings at low bed densities. New For 7:63–92
South DB (2000) Planting morphologically improved pine seedlings to increase survival and growth. Ala.
Agric. Exp. Sta. Auburn Univ. For. and Wildlife Res. Ser. No. 1
South DB, Barnett JP (1986) Herbicides and planting affect early performance of container-grown and bare-
root loblolly pine seedlings in Alabama. New For 1:17–27
South DB, Blake JI (1994) Top-pruning increases survival of pine seedlings. Ala Agric Exp Sta Highlights
Agric Res 41:9
South DB, Donald DGM (2002) Effect of nursery conditioning treatments and fall fertilization on survival
and early growth of Pinus taeda seedlings in Alabama, U.S.A. Can J For Res 32:1171–1179
South DB, Hallgren SW (1997) Research versus operational correlations between seedling survival and root
growth potential of shortleaf pine. New For 13:357–365
South DB, Mexal JG (1984) Growing the ‘‘best’’ seedling for reforestation success. Ala Agric Exp Stn
Auburn Univ For Dep Ser No. 12
South DB, Mitchell RJ (1999) Determining the ‘‘optimum’’ slash pine seedling size for use with four levels
of vegetative management on flatwoods site in Georgia, USA. Can J For Res 29:1039–1046
South DB, Stumpff NJ (1990) Root stripping reduces root growth potential of loblolly pine seedlings. South
J Appl For 14:196–199
South DB, Boyer JN, Bosch L (1985) Survival and growth of loblolly pine as influenced by seedling grade:
13 year results. South J Appl For 9:76–81
South DB, Rakestraw JL, Lowerts GA (2001) Early gains from planting large diameter seedlings and
intensive management are additive for loblolly pine. New For 21:97–110
New Forests (2012) 43:711–738 735
123
South DB, Harris SW, Barnett JP, Hainds MJ, Gjerstad DH (2005) Effect of container type and seedling size
on survival and early height growth of Pinus palustris seedlings in Alabama, U.S.A. For Ecol Manage
204:385–398
Stewart JD, Bernier PY (1995) Gas exchange and water relations of 3 sizes of containerized Picea mariana
seedlings subjected to atmospheric and edaphic water stress under controlled conditions. Ann For Sci
52:1–9
Stone EC (1955) Poor survival and the physiological condition of planting stock. For Sci 1:89–94
Stone EC, Cavallaro JI, Norberg EA (2003) Critical RGC-Expected survival models for predicting survival
of planted white fir (Abies concolor Lindl.) seedlings. New For 26:65–82
Stupendick JAT, Shepherd KR (1980) Root regeneration of root-pruned Pinus radiata seedlings. II. Effects
of root-pruning on photosynthesis and translocation. N Z J For Sci 10:148–158
Sullivan TP, Moses RA (1986) Demographic and feeding responses of a snowshoe hare population to habitat
alteration. J Appl Ecol 23:53–63
Sutton RF (1980) Planting stock quality, root growth capacity, and field performance of three boreal
conifers. N Z J For Sci 10:54–71
Sutton RF (1990) Root growth capacity in coniferous forest trees. Hortic Sci 25:259–266
Sword Sayer MA, Sung SJ, Haywood JD (2011) Longleaf pine root system development and seedling
quality in response to copper root pruning and cavity size. South J Appl For 35:5–11
Tan W (2007) Impacts of nursery cultural treatments on stress tolerance in 1?0 container white spruce
(Picea glauca [Moench] Voss) seedlings for summer-planting. New For 33:93–107
Tan W, Blanton S, Bielech JP (2008) Summer planting performance of white spruce 1?0 container seed-
lings affected by nursery short-day treatment. New For 35:187–205
Tanaka Y, Walstad JD, Borrecco JE (1976) The effect of wrenching on morphology and field performance
of Douglas-fir and loblolly pine seedlings. Can J For Res 6:453–458
Teskey RO, Hinckley TM (1986) Moisture: effects of water stress on trees. In: Hennessey TC, Dougherty
PD, Kossuth SV, Johnson JD (eds) Stress physiology and forest productivity. Martinus Nijhoff Pub-
lishers, The Netherlands, pp 9–33
Thiffault N (2004) Stock type in intensive silviculture: a (short) discussion about roots and size. For Chron
80:463–468
Thomas DS (2009) Survival and growth of drought hardened Eucalyptus pilularis Sm. seedlings and
vegetative cuttings. New For 38:245–259
Thompson BE (1985) Seedling morphological evaluation: what you can tell by looking. In: Duryea ML (ed)
Evaluating seedling quality: principles, procedures, and predictive ability of major tests. Forest
Research Laboratory, Oregon State University, Corvallis, OR, pp 59–72
Tillotson CR (1915) Forest planting in the eastern United States. USDA Bull. 153
Timmer VR (1997) Exponential nutrient loading: a new fertilization technique to improve seedling
performance on competitive sites. New For 13:279–299
Timmer VR, Aidelbaum AS (1996) Manual for exponential nutrient loading of seedlings to improve
outplanting performance on competitive forest sites. NODA/NFP Tech. Rep. TR-25
Timmer VR, Miller BD (1991) Effects of contrasting fertilization and irrigation regimes on biomass,
nutrients, and water relations of container grown red pine seedlings. New For 5:335–348
Timmer VR, Armstrong G, Miller BD (1991) Steady-state nutrient preconditioning and early out-planting
performance of containerized black spruce seedlings. Can J For Res 21:585–594
Timmis R, Tanaka Y (1976) Effects of container density and plant water stress on growth and cold hardiness
of Douglas-fir seedlings. For Sci 22:167–172
Tinus RW (1974) Characteristics of seedlings with high survival potential. In: Tinus RW, Stein WI, Balmer
WE (eds) Proceedings of the North American containerized forest tree seedling symposium. Great
Plains Ag. Council Publ. No. 68, pp 276–282
Tinus RW, McDonald SE (1979) How to grow tree seedlings in containers in greenhouses. USDA Forest
Service Gen. Tech. Rep. RM-60
Toumey JW (1916) Seeding and planting. Wiley, New York
Troth JL, Campbell RG, Allen HL (1986) Nutrients: use of forest fertilization and nutrient efficient
genotypes to manage nutrient stress in conifer stands. In: Hennessey TC, Dougherty PD, Kossuth SV,
Johnson JD (eds) Stress physiology and forest productivity. Martinus Nijhoff Publishers, The Neth-
erlands, pp 61–99
Tsakaldimi MN, Ganatsas PP (2006) Effects of chemical pruning on stem growth, root morphology and field
performance on the Mediterranean pine Pinus halepensis Mill. Sci Hortic 109:183–189
Tuttle CS, South DB, Golden MS, Meldahal RS (1987) Relationship between initial seedling height and
survival and growth of loblolly pine seedlings planted during a droughty year. South J Appl For
11:139–143
736 New Forests (2012) 43:711–738
123
Tuttle CS, South DB, Golden MS, Meldahal RS (1988) Initial Pinus taeda seedling height relationships with
early survival and growth. Can J For Res 18:867–871
Unterschuetz PF, Ruetz W, Geppert R, Ferrell WK (1974) The effect of age, pre-conditioning, and water
stress in the transpiration rates of Douglas-fir (Pseudotsuga menziesii) seedlings of several ecotypes.
Physiol Plant 32:214–221
Vaartaja O (1960) Effect of photoperiod on drought resistance of white spruce seedlings. Can J Bot
38:597–599
van den Driessche R (1969) Influence of moisture supply, temperature, and light on frost hardiness changes
in Douglas-fir seedlings. Can J Bot 47:1765–1772
van den Driessche R (1980) Effects of nitrogen and phosphorus fertilization on Douglas-fir nursery growth
and survival after outplanting. Can J For Res 10:65–70
van den Driessche R (1984) Relationship between spacing and nitrogen fertilization of seedlings in the
nursery, seedling mineral nutrition, and outplanting performance. Can J For Res 14:431–436
van den Driessche R (1985) Late-season fertilization, mineral nutrient reserves, and retranslocation in
planted Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. For Sci 31:485–496
van den Driessche R (1988) Nursery growth of conifer seedlings using fertilizers of different solubilities and
application time, and their forest growth. Can J For Res 18:172–180
van den Driessche R (1991a) Influence of container nursery regimes on drought resistance of seedlings
following planting: survival and growth. Can J For Res 21:555–565
van den Driessche R (1991b) Effects of nutrients on stock performance in the forest. In: van den Driessche R
(ed) Mineral nutrition of conifer seedlings. CRC Press, Boca Raton, FL, pp 229–260
van den Driessche R (1992) Changes in drought resistance and root growth capacity of container seedlings
in response to nursery drought, nitrogen, and potassium treatments. Can J For Res 22:740–749
VanderSchaaf C, McNabb K (2004) Winter nitrogen fertilization of loblolly pine seedlings. Plant Soil
265:295–299
Verdauger D, Vilagran J, Lloansi S, Fleck I (2011) Morphological and physiological acclimation of Quercus
coccifera L. seedlings to water availability and growing medium. New For 42:363–381
Villar-Salvador P, Ocana L, Penuelas J, Carrasco I (1999) Effects of water stress conditioning on the water
relations, root growth capacity, and nitrogen and non-structural carbohydrate concentration of Pinus
halepensis Mill. seedlings. Ann For Sci 56:459–465
Villar-Salvador P, Penuelles R, Enriquez E, Penuelas J, Rubira JL (2004a) Nursery cultivation regimes,
plant functional attributes, and field performance relationship in Mediterranean oak Quercus ilex L. For
Ecol Manage 196:257–266
Villar-Salvador P, Penuelles R, Oliet J, Enriquez E, Penuelas JL, Jacobs DF, Gonzalez M (2004b) Drought
tolerance and transplanting performance of holm oak (Quercus ilex) seedlings after drought hardening
in the nursery. Tree Physiol 24:1147–1155
Wagner B, Colombo SJ (eds) (2001) Regenerating Ontario’s forests. Fitzhenry & Whiteside Ltd., Markham, ON
Wakeley PC (1948) Physiological grades of southern pine nursery stock. Soc Am For Proc 1948:311–322
Wakeley PC (1954) Planting the southern pines. Agric. Monogr. No. 18. USDA Forest Service Washington,
DC, 233 pp
Webb RA (1972) Use of boundary line in the analysis of biological data. J Hortic Sci 97:309–319
Wenny DL, Liu Y, Dumroese RK, Osborne HL (1988) First year field growth of chemically root pruned
containerized seedlings. New For 2:111–118
Whitcomb CE (1984) Plant production in containers. Lacebark Publications, Stillwater, OK
Williams HM, South DB (1992) Effects of fall fertilizer applications on mitotic index and bud dormancy of
loblolly pine seedlings. For Sci 38:336–349
Williams BJ, Pellett NE, Klein RM (1972) Phytochrome control of growth cessation and initiation of cold
acclimation in selected woody plants. Plant Physiol 50:262–265
Williams HM, South DB, Webb A (1988) Effects of fall irrigation on morphology and root growth potential
of loblolly pine seedlings growing in sand. S Afr For J 147:1–5
Wilson BC, Jacobs DF (2006) Quality assessment of temperate zone deciduous hardwood seedlings. New
For 31:417–433
Wilson ER, Vitols K, Park A (2007) Root characteristics and growth potential of container and bare-root
seedlings of red oak (Quercus rubra L.) in Ontario Canada. New For 34:163–176
Young LJ (1921) Forest planting in southern Michigan. J For 19:1–8
Young E, Hanover JW (1978) Effects of temperature, nutrient, and moisture stresses on dormancy of blue
spruce seedlings under continuous light. For Sci 24:458–467
Zida D, Tigabu M, Sawadogo L, Ode
´n PC (2008) Initial seedling morphological characteristics and field
performance of two Sudanian savannah species in relation to nursery production period and watering
regimes. For Ecol Manage 255:2151–2162
New Forests (2012) 43:711–738 737
123
Zobel BJ, Talbert JT (1984) Applied forest tree improvement. Wiley, New York
Zwiazek JJ, Blake TJ (1989) Effects of preconditioning on subsequent water relations, stomatal sensitivity,
and photosynthesis in osmotically stressed black spruce. Can J Bot 67:2240–2244
Zwolinski JB, South DB, Cunningham L, Christie S (1996) Weed control and large bare-root stock improve
early growth of Pinus radiata in South Africa. N Z J For Sci 26:163–172
738 New Forests (2012) 43:711–738
123
