New Phytol.

(1998), 139, 505–516

Germination and post-germination growth

to carrot seedling emergence : predictive
threshold models and sources of variation
between sowing occasions

B  W. E. F I N C H-S A V A G E*, J. R. A. S T E C K E L    K. P H E L P S
Horticulture Research International, Wellesbourne, Warwick CV35 9EF, UK
(Received 11 November 1997 ; accepted 19 March 1998)


Cumulative germination curves were recorded for carrot (Daucus carota L.) seeds at a range of constant
temperatures (T ) and water potentials (Ψ) in the laboratory and under variable soil conditions in 15 seed-bed
environments in the field. A single base temperature (Tb), a distribution of base water potentials (Ψb(G)) for
percentiles (G) of the population and the hydrothermal time constant (Hg) were determined from laboratory data.
Although less effective at low Ψ, it was possible, using these germination parameters, to satisfactorily describe the
effect of T and Ψ on germination rates under constant conditions according to the threshold models of thermal
and hydrothermal time. These models were applied to field data with the condition that the germination process
ceased if T ! Tb for thermal time and additionally Ψ ! Ψb(G) for hydrothermal time.
Neither model accurately predicted germination patterns in the field. However, the pattern of germination was
adequately described in most situations by a modified threshold model in which the predicted progress towards
germination was unaffected by soil Ψ, provided it remained above Ψb(G), and was therefore more rapid under
variable seed bed conditions than hydrothermal time. In this modified threshold model, the condition Ψ " Ψb(G)
had to be fulfilled at the initiation of radicle extension before germination occurred. This result implies that the
initiation of radicle growth operates as a moisture-sensitive step that can determine germination and seedling
emergence timing under variable soil-moisture conditions.
Seedling emergence was also recorded in the field and used to determine, separately, the impact of germination
and post-germination growth on the variation in seedling emergence patterns. The analysis suggests that delays
in seedling emergence occur largely in the germination phase, but that seedling losses and variation in the spread
of seedling emergence times within the population occur largely during the post-germination growth phase.

Key words : Carrot (Daucus carota L.), germination, seedling emergence, crop establishment, threshold models,
thermal time, hydrothermal time.

therefore directly influences both their yield and


The pattern of seedling emergence within a popu-
lation is determined by a complex interaction of
ambient weather conditions, soil, seed and seedling
characteristics. These interactions are little under-
stood in the natural environment. However, some
progress is being made in the modelling and
prediction of seedling emergence patterns in hor-
ticulture. Here the situation is less complex as soil is
prepared uniformly, and the whole population of
seeds (usually non-dormant) is sown at the same
time and to a uniform depth. Despite this, seedling
emergence in vegetable crops is still variable and
* To whom correspondence should be addressed.
E-mail : bill.finch-savage!hri.ac.uk
monetary value (Finch-Savage, 1995). For example,
in carrots, plant density determines both total yield
and mean root size (Bleasdale, 1967) and the
uniformity of seedling emergence times can greatly
influence the uniformity of plant size at harvest and
therefore graded yields (Benjamin, 1982). An under-
standing of the causes of variation in seedling
emergence both within and between sowing
occasions is therefore necessary to determine efficient
crop production practices.
The pre-germination and post-germination phases
of the seedling emergence period are each uniquely
affected by adverse seed-bed conditions (Finch-
Savage, 1995). Although the timing of germination
in the seed bed can account for much of the variation
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506 W. E. Finch-Savage, J. R. A. Steckel and K. Phelps

in the timing of onion seedling emergence (Finch- mated from controlled conditions, could describe
Savage & Phelps, 1993) it is thought to be the post- germination patterns that occur under the variable
germination phase where most pre-emergence seed- conditions experienced following sowing into hor-
ling losses occur (Hegarty & Royle, 1978 ; Durrant, ticultural seed beds.
1981). However, information on the relative con-
tribution of the pre-germination and post-germi-   
nation phases to the variation in seedling emergence
both within and between sowing occasions is very Field experiment
limited.
The experiment was sown in 1988 at Wellesbourne,
Under more extreme conditions, seed-bed factors
UK into a sandy loam soil of the Wick series
such as increased soil impedance and reduced oxygen
(Whitfield, 1974) and had a split-plot design with
supply can have an overriding influence on seedling
three replicates. Main plots were five sowing
emergence. However, under good horticultural
occasions (30 Mar. ; 8 Apr. ; 6 May ; 3 June ; 3 Aug.)
practice, soil temperature and water potential are
and subplots were three irrigation treatments. Ir-
arguably the major influences on the timing and
rigation treatments were : no irrigation ; 12±5 mm
pattern of seed germination and seedling emergence
applied more than 90 °C d (" 1 °C) after sowing
in the field. The threshold models of thermal time
(pre-emergence) ; 12±5 mm applied 48 h before
(Bierhuizen & Wagenvoort, 1974 ; Garcia-Huidobro,
sowing (pre-sowing) plus pre-emergence. Each sub-
Monteith & Squire, 1982) and hydrothermal time
plot was a single 1±5-m row containing 100 carrot
(Gummerson, 1986 ; Dahal & Bradford, 1994 ;
(Daucus carota L. ; cv. Nantura) seeds sown 15-mm
Bradford, 1995) have been developed to describe the
deep with a hand-pushed cone seeder (Craftsman
germination responses of a population of seeds to
Machine Co., Winnipeg, Canada). All plots were
temperature and water potential under constant
sprayed with a soil conditioner (Soiltex2 L1, Inter-
conditions, but little is known of their descriptive
lates Ltd., Skelmersdale, UK, at 125 l ha−" in 1000 l
ability under field conditions.
of water) following sowing. Pests and diseases were
Seed advancement studies which limit moisture
controlled as in good commercial practice, and soil
availability to seeds (i.e. osmotic priming) have
samples, for moisture determinations, were taken at
shown that metabolic advancement occurs at levels
sowing depth in all plots at sowing time.
of water stress which prevent radicle emergence
In plots adjacent (300-mm apart) to those sown
from the seed (Khan, 1992 ; Bradford & Haigh,
with seeds, identical plots were sown with 9, 50-mm
1994). In most species radicle extension growth is
lengths of dry Visking2 tube (6±3-mm diameter,
also less sensitive to moisture stress than radicle
Medicell International Ltd, London, UK) each
emergence from the seed (Ross & Hegarty, 1979).
containing 50 seeds. At regular intervals following
Taken together, this suggests that the initiation of
sowing a tube was exhumed from each replicate plot
radicle growth is a moisture-sensitive, rate-limiting
and the number of germinated seeds were counted.
step in the progress of seedling emergence from the
The tubes were perforated at 5-mm intervals along
soil. The interaction of this step and variable soil
their full length and the ends folded and stapled. In
moisture levels can largely determine the pattern of
this form the tube acted merely as an easily
onion seedling emergence in the field (Finch-Savage
recoverable container for seeds which had good soil
& Phelps, 1993). This suggests that additional
contact, and would wet and dry with the seed bed. In
conditions might need to be applied to the ac-
preliminary tests in soil-filled trays the timing of
cumulation of time in threshold models to take
germination in these tubes accurately represented
account of the influence of this moisture-sensitive
that of seeds sown directly into the soil.
step in the prediction of germination and therefore
Seedling emergence and germination in the field
seedling emergence patterns under field conditions.
were recorded at regular intervals. From these
Although experiments under controlled conditions
recordings, times to germination or emergence of
suggest that the post-germination growth which
different percentiles of the seed population were
leads to seedling emergence is also quantifiable using
calculated in thermal time (°C d) above the base
threshold models (Fyfield & Gregory, 1989 ; Wheeler
temperature determined in laboratory experiments
& Ellis, 1991) few data are available where this phase
(2±15 °C). Procedures for this are described later.
of seedling emergence in the field is not confounded
Final percentage germination and emergence were
with germination.
also calculated and subjected to angular transform-
In the present work an attempt was made to study
ation. All germination or seedling emergence
the influence of seed-bed conditions on both pre-
variables were then subjected to split-plot .
germination and post-germination phases of carrot
seedling emergence to assess their relative con-
Constant environment experiments
tribution to variation in seedling emergence patterns.
A further objective was to determine whether Germination in four replicates of 50 carrot seeds (D.
existing threshold models, using parameters esti- carota ; cv. Nandor) was recorded on moist absorbent

Carrot germination and post-germination growth
paper (Whatman2, grade 181) at c. 5 °C intervals
between 5 and 35 °C and at a range of six water
potentials (ψ) between 0±0 and ®1±2 MPa at 20 °C.
Temperature was monitored continuously by
thermistors placed, like seeds, on moist paper. Water
potentials were established using polyethylene glycol
(PEG, 8000, BDH Ltd, Poole, UK) solutions made
up according to Michel (1983). The same solution
volume : filter paper ratio was used for germination
as that in the vapour pressure osmometer (model
5100C ; Wescor Inc, Logan, UT, USA) used to
measure water potential. Thus the problem of filter-
paper exclusion of PEG pointed out by Hardegree &
Emmerich (1990) was avoided.
For comparison with field data, seed germination
and seedling emergence were determined in the
absence of moisture stress at 20 °C in seedling trays
filled with soil taken from the field experiment site.
The soil was sieved and dried before water was
added to bring the gravimetric soil moisture content
to 12±5 % which was the equilibrium value at
0±005 MPa (nominal field capacity) determined from
water release curves. For emergence counts, 50 seeds
were sown in rows on a level soil surface and covered
with 15 mm of soil from the same batch. For
germination 9, 50-mm lengths of dry Visking tube
(as sown in the field) each containing 50 seeds were
similarly sown. The soil was firmed by applying a
weight approx. equal to that of the rear wheel of the
seeder used in the field experiment. Germination
and seedling emergence were recorded under
fluorescent lights in six replicate trays placed in
plastic bags to minimize moisture loss.
Analysis of data
Germination models. Threshold germination models
used in this work are based on the theories of thermal
and hydrothermal time of Garcia-Huidobro et al.
(1982) and Gummerson (1986) respectively. It is an
assumption of these theories that seeds germinate in
a set order and that this order is unaffected by
germination conditions. Each seed can then be
assigned a value of G, which is the fraction of the
population at which it germinates. As the ger-
mination rate of fraction G was linearly related to
sub-optimal temperatures (T ), the rate of germi-
nation of that fraction G was defined by Gummerson
(1986) as :
1}t(G) ¯ (T®Tb)}θ(G), (1)
where t is the time to germination and θ the thermal
time to germination. The base temperature (Tb(G))
below which germination does not occur was defined
as the intercept on the temperature axis. According
to the theory of hydrothermal time (Gummerson,
1986) rates in thermal time are proportional to water
potential thus :
1}θ(G) ¯ (Ψ®Ψb(G))}Hg, (2)
14698137, 1998, 3, Downloaded from https://nph.onlinelibrary.wiley.com/doi/10.1046/j.1469-8137.1998.00208.x by CAPES, Wiley Online Library on [08/08/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
507
where Hg (hydrothermal time) is a constant and, as
there was a linear relationship between the ger-
mination rate of fraction G and Ψ, the base water
potential (Ψb(G)) below which germination did not
occur was defined as the intercept on the Ψ axis.
Substitution of (2) in (1) gives :
1}t(G) ¯ (T®Tb) (Ψ®Ψb(G))}Hg. (3)
From (3) it can be seen that the distribution of times
to germination are influenced by the variability of
Ψb(G).
Hydrothermal time has been used to describe
germination curves produced in a wide range of
combinations of constant temperature and water
potential (Gummerson, 1986 ; Dahal & Bradford,
1994). Under field conditions both temperature and
soil water potential vary unpredictably. However, it
was argued by Finch-Savage and Phelps (1993) that
under good horticultural practice seeds are sown into
moist soil so that initial imbibition is rapid. Once
imbibed, the seed would then progress towards
germination unaffected by Ψ provided it remained
above Ψb(G). Thus eqn (3) would become :
1}t(G) ¯ (T®Tb) (®Ψb(G))}Hg. (4)
Parameter estimation. The techniques for parameter
estimation were analogous to those used in thermal
time (Garcia-Huidobro et al., 1982). The parameters
of eqn (1), Tb and θ(50) were estimated from data
collected at 0 MPa by linear regression of 1}t(50)
against sub-optimal temperatures. θ(10), θ(20) etc.
were estimated by interpolation of germination
curves from all temperatures on the thermal time
scale. Data from all water potentials (0 to ®1±2 MPa)
at 20 °C were used to estimate Ψb(10), Ψb(20) etc.
The values of Tb and Ψb(50) were substituted in eqn
(3) and Hg was estimated by non-linear regression
using 1}t(50) values from all controlled environment
regimes.
Application of models to field data. Using recorded
soil moistures as initial values, a model, developed
for these soils (Walker & Barnes, 1981), was used to
estimate soil moisture and temperature at sowing
depth at 6-h intervals following sowing. The model
required daily air temperature and rainfall measure-
ments made at an agrometeorological station within
0±5 km of the plots. A water release curve was
determined, using pressure membrane apparatus, and
used to convert estimates of soil moisture produced
by the model to soil water potential.
The ability of eqns (1), (3) and (4) to describe seed
germination recorded under variable field conditions
was then tested. To use these equations, germination
for any fraction G was assumed to occur after n 6-h
periods where n was defined as laid out below and
T(i), Ψ(i) are the temperature and water potential for
each period. The conditions for germination for each
equation then become as follows.
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508 W. E. Finch-Savage, J. R. A. Steckel and K. Phelps

Eqn (1) : considered the ‘ best ’ estimate of Tb. Significant

negative linear relationships (r " 0±97, P ! 0±01)
n
θ(G) & 0±25 3 (T(i)®Tb) T(i) " Tb. were also found between the germination rate of
i=! deciles 10–80 % of the seed population and Ψ in the
Eqn (3) : range 0±0 and ®0±8 MPa at 20 °C (Fig. 2). According
to eqn (2), lines corresponding to deciles should be
n parallel, but the variability between deciles at Ψ ¯
Hg & 0±25 3 (T(i)®Tb) (ψ(i)®ψb(G))
i=! 0 MPa was considerably more than that at Ψ ¯
T(i) " Tb, ψ(i) " ψb(G). 0±8 MPa and so it was not appropriate to fit parallel
lines. Therefore, Ψb(G) were estimated as the
Eqn (4) : intercepts of a set of converging lines. The resulting
n estimates were well described by the following
Hg & 0±25 3 (T(i)®Tb) (0®ψb(G)) equation :
i=!
T(i) " Tb, ψ(i) " ψb(G). Ψb(G) ¯®0±9745®0±0010145 G­0±000040782 G#
(R# " 0±98, P ! 0±001 ; Fig. 2 insert).

Hydrothermal time (Hg) estimated from eqn (2)
was 47±71 MPa °C d. Using the estimates of Tb, the
Constant environment experiments
values of Ψb(G) and Hg all germination time courses
Significant positive linear relationships (r " 0±98, were plotted in hydrothermal time (Fig. 3). With the
P ! 0±01) were found between the germination rate exception of data from ®0±8 MPa and deciles greater
on water (0±0 MPa) of deciles 10–80 % of the seed than 60 at ®0±6 MPa germination curves collapsed
population and temperature up to 25 °C (Fig. 1). onto a single line. Mean final percentage germination
The intercept on the temperature axis of the 50th was not significantly different at water potentials of
percentile was taken as the base temperature (Tb ¯ ®0±6 MPa or greater. However, final percentage
2±15 °C). There was a difference in the intercepts of germination was reduced at ®0±8 MPa (Fig. 3 ;
other percentiles to give a  of 0±178 °C. The 50th P ! 0±01) and no germination was recorded at
percentile, as median of the distribution, was ®1±2 MPa.

0·7 _, 10 %
^, 20 %
+, 30 %
*, 40 %
0·6 E, 50 %
D, 60 %
y, 70 %
x, 80 %
0·5
Rate of germination (1/t(G))

0·4

0·3

0·2

0·1

0·0
0 2 4 6 8 10 12 14 16 18 20 22 24
Temperature °C

Figure 1. The effect of constant temperature on the rate of germination for deciles of the Daucus carota seed
population. Extrapolation of the linear regression fitted to data for the 50th percentile to the temperature axis
(Tb) is shown.
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Carrot germination and post-germination growth 509

100 0·6

80

Percentile
60
0·5
40
20
0

Rate of germination (1/t(G))

–1·0 –0·9 –0·8 –0·7 0·4
¾b(G) MPa

_, 10 %
^, 20 % 0·3
+, 30 %
*, 40 %
E, 50 %
D, 60 %
y, 70 % 0·2
x, 80 %

0·1

0·0
–1·2 –1·0 –0·8 –0·6 –0·4 –0·2
Water potential (MPa)
Figure 2. The effect of constant water potential on the rate of germination for deciles of the seed population.
The inset shows the distribution of Ψbs (intercepts with the water potential axis) for different deciles of the
population, a quadratic regression (see text) is fitted to the data.

100

80

Temperature
E, 5 °C
Germination (%)

60 +, 10 °C
_, 15 °C
y, 20 °C
U, 25 °C
Water potenial
40 D, – 0·0 MPa
*, – 0·2 MPa
^, – 0·4 MPa
x, – 0·6 MPa
V, – 0·8 MPa
20

0
0 50 100 150 200
Hydrothermal time (MPa °Cd)
Figure 3. The hydrothermal time courses of germination of Daucus carota seeds at a range of constant
temperatures and water potentials.

following sowing into three irrigation treatments on

Field experiments
five sowing occasions. Mean temperatures at sowing
Application of germination models to field data. Figure depth during germination were 10±2, 11±1, 15±0, 17±6,
4 shows cumulative germination curves recorded 21±1 °C for sowings 1–5 respectively. Soil moisture
 510

No irrigation Pre-emergence irrigation Pre-sowing + pre-emergence irrigation

100
(a) (b) (c)
75
50 Sowing 1

25
0
100
(d) (e) (f )
75
50
Sowing 2

25
0
100
(g) (h) (i )
75
50
Sowing 3

25
0
100
(j) (k) (l )
W. E. Finch-Savage, J. R. A. Steckel and K. Phelps

75
50
Sowing 4

25
0
100 (m) (n) (o)
75
50
Sowing 5

25
0
0 5 10 15 20 25 30 35 0 5 10 15 20 25 30 35 0 5 10 15 20 25 30 35

Figure 4. The time courses of germination (——) in 15 different seed-bed environments. Predictions of germination time for each
decile of the population according to eqns (1) (E), (3) (+) and (4) (*) subject to the conditions outlined in the text are also shown.
Sowings 1–5 are shown vertically e.g. (a), (d ), (g), ( j), (m) respectively. Irrigation treatments are shown horizontally e.g. no
irrigation (a), pre-emergence (b), pre-sowing­pre-emergence (c).

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Carrot germination and post-germination growth
100
90
80
Percentage germination
70
60
50
40
30
20
10
0 sowings 3 and 5 where prolonged dry periods
a b c a b c a b c a b c a
1 2 3 4
Sowing
Figure 5. Percentage germination (*) and percentage
seedling emergence (8 9) in 15 different seedbed environ-
ments. At each sowing the columns from left to right are :
a, no irrigation ; b, pre-emergence irrigation ; c, pre-
sowing­pre-emergence irrigation treatment respectively.
Horizontal lines are germination (——) and seedling
emergence (– – – – –) under non-stressed conditions.
patterns were very different with prolonged moist
periods following sowing 2 and prolonged dry
periods following sowings 3 and 5, conditions were
more variable following sowings 1 and 4. Under
these differing conditions, rate and spread of ger-
mination varied greatly between the fifteen different
seed-bed environments, but final percentage ger-
mination was consistently high (Fig. 4). These
germination characteristics are further described in
the next section.
Figure 4 shows germination time simulations from
eqns (1), (3) and (4) plotted alongside recorded
germination data in real time following sowing. With
the exception of the relatively moist environment
experienced at sowing 2, thermal time (eqn (1))
underestimated the time required for germination.
Hydrothermal time (eqn (3)) with the exception of
some deciles at sowing 4 greatly overestimated the
time required for germination. With the exception of
sowing 2 the pattern of germination is generally best
described by eqn (4).
Sources of variation in seedling emergence. Final
percentage germination was little affected by seed-
bed environment (sowing and irrigation treatment)
with mean germination across seed-bed environ-
ments of 94 % (Fig. 5). However, there was a
significant reduction (P ! 0±001) in final percentage
seedling emergence at later sowing dates. These data
suggest that the cause of variation in percentage
seedling emergence occurs largely post-germination.
Presenting the pattern of germination and emerg-
ence in thermal time allows the more consistent
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511
seasonal effect of temperature to be eliminated to
show the impact of other seed-bed factors. Thermal
time to germination and the thermal time required
for post-germination growth to seedling emergence
(emergence time–germination time) for deciles
(10–80) of the population are therefore shown in
Figure 6.
The degree of delay caused by seed-bed stress in
each of the two phases of seedling emergence can be
seen by comparison with curves that were de-
termined under unstressed conditions in seed trays
filled with moist soil (Fig. 6). It is clear that variation
in seedling emergence time results from delays in
both the pre-germination and post-germination
phases. Delay in germination time was greatest in
b c
5
occurred. This effect was less marked in the time
required for post-germination growth to emergence.
In addition, delays in germination time were not
necessarily associated with slow post-germination
growth. However, significant delays in germination
time were associated with reduced percentage seed-
ling emergence (Fig. 5).
In two seed-bed environments the thermal time
required for post-germination growth to emergence
was less than that recorded under unstressed con-
ditions. The reasons for this are not known, but this
situation could have resulted from soil slumping or
erosion following rain and}or irrigation which would
effectively reduce sowing depth.
In the absence of seed-bed stress, post-germi-
nation growth rate was similar for all percentiles of
the population, but tended to decrease at higher
percentiles. As seed-bed stress increased (greater
delay in thermal time) post-germination growth rate
of higher percentiles (the last to germinate) decreased
more than lower percentiles and this increased the
spread of time of post-germination growth to
seedling emergence in the population. This effect
was much less marked in the germination phase.
Thus most variation in the spread of seedling
emergence times in the population between seed-bed
environments appears to occur post-germination.

Threshold germination models under constant
conditions
There is currently much interest in the use of
threshold models to explain and describe the ger-
mination response of seeds to a range of constant
stimuli (Bradford, Dahal & Ni, 1993). These models
are increasingly being used as a framework in which
the mechanisms of germination can be understood.
However, they are derived from, and usually tested
against, data collected under constant conditions and
therefore their true descriptive abilities are not
known.
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512 W. E. Finch-Savage, J. R. A. Steckel and K. Phelps

90
(a)
80

70

60

50

40

30

20

10
Percentile

90
(b)
80

70

60

50 Sowing time
D, 1
40 E, 2
*, 3
30 +, 4
^, 5
20

10

0
0 50 100 150 200 250 300

Thermal time (°C d)

Figure 6. Thermal time required for (a) germination and (b) post-germination growth (emergence –
germination) to emergence. , non-stressed conditions. Irrigation treatment : ——, no irrigation ; – – –, pre-
emergence ; [[[[[[, pre-sowing­pre-emergence.

In the present work, although less effective at
lower water potentials, it was possible to describe the
germination response of carrot seeds at a range of
constant temperatures and water potentials using, a
single Tb, a distribution of Ψb(G) and Hg according to
the principles of hydrothermal time (eqn (3) ; Fig. 3).
There was variation in the estimate of Tb between
deciles of the population similar to that shown by
Marshall & Squire (1996). Here a single Tb has been
used, as this is the currently accepted practice when
calculating hydrothermal time, but an alternative
approach has been suggested elsewhere (Phelps &
Finch-Savage, 1997).
There can be subtle differences in Ψb between
tomato genotypes (Dahal & Bradford, 1990) and
θ(50) also varies, but it has been suggested that Tb
might be a species characteristic and vary little
between genotypes (Ellis et al., 1986 ; Dahal,
Bradford & Jones, 1990). Clearly germination rates
differ between cultivars (Corbineau, Picard & Co# me,
1994), but it is likely that in carrots, bases do not
vary greatly as a very similar Ψb (®0±81 MPa ; Ross
& Hegarty, 1979) and a similar Tb (1±4 °C ; Bierhuizen
& Wagenvoort, 1974) were recorded for cvs
unrelated to those used here. In the experiments
presented, a seed lot from a closely related genotype
was selected for constant environment experiments
(cv. Nandor) that had germination characteristics
similar to the seed lot used in field experiments (cv.
Nantura). Estimates of Tb (1±6³0±364 °C) and θ(50)
(58 °C d) for ‘ Nantura ’ from limited constant-
environment data were very similar to those de-
termined for ‘ Nandor ’ (2±15³0±178 °C and 56±5 °C d
respectively). Similar spreads in thermal time (θG
10–80 % for ‘ Nantura ’ 40 °C d and Nandor 36 °C d)
show that a very similar range of Ψbs exist in both
seed lots according to the theory of hydrothermal
time. In relation to the variation that exists even

Carrot germination and post-germination growth
within a good seed bed these differences are subtle
and had little affect on field predictions.
Threshold germination models under field conditions
Neither the thermal (eqn (1) nor hydrothermal (eqn
(3)) time models adequately described patterns of
carrot germination in the field. Thermal time
underestimated, and hydrothermal time greatly
overestimated, the time required for germination.
Clearly, when applying these models to a field
situation there are many assumptions and
inaccuracies, not least the estimation of soil Ψ from
meterological data, which might have led to this lack
of fit. However, the pattern of germination was
adequately described in most situations by a
modified hydrothermal time model in which the
predicted progress towards germination was more
rapid as it was unaffected by soil Ψ provided it
remained above Ψb(G) (eqn (4)). Further consider-
ation of the results is therefore justified.
A straightforward thermal time approach was
unlikely to be effective because the soil at 15 mm
(sowing depth) dries very quickly in the absence of
rain to delay germination in thermal time. The
inadequate description of the data by hydrothermal
time and the improved fit using eqn 4 is, however,
more difficult to explain.
When calculating hydrothermal time it is assumed
that Tb, Ψb(G) and Hg are intrinsic properties of the
seed which remain the same under all germination
conditions. However, experiments under constant
conditions have shown that there are examples of
interactions, which might be species-specific, be-
tween Ψb(G) and T and between Ψb(G) and Ψ
(Dahal & Bradford, 1994). Both Gummerson (1986)
and Dahal & Bradford (1994) and the present work
show that hydrothermal time is less descriptive of
germination time courses as Ψ decreases. This is
because seeds appear to adapt physiologically by
increasing osmotic potential and lowering Ψb(G)
when exposed for prolonged periods to low Ψ (Ni
and Bradford, 1992). This accounts for much of the
inaccuracy in the use of hydrothermal time under
constant conditions (Dahal & Bradford, 1994).
Under these conditions an alternative model based
on the nominal accumulation of osmotically-active
compounds (Rowse, Goodman & Grey, 1986) might
be more descriptive. However, under good hor-
ticultural practice with effective use of irrigation
(Finch-Savage, 1990 a, b) such prolonged periods of
low Ψ should not occur in the seed bed. In the
current work, these conditions might have occurred
at sowings 3 and 5 where irrigation was not applied
(Fig. 4 g, m).
A further implicit assumption of hydrothermal
time is that seeds wet up and dry as rapidly, and to
the same extent, as the surrounding soil and that the
impact of this on the rate of progress towards
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513
germination, including radicle emergence, is equal
throughout the germination process. There is no
compelling reason to believe this is the case. Indeed
Finch-Savage & Phelps (1993) based on the work of
Hegarty & Ross (1978) and Ross & Hegarty (1979)
suggest that in many species the initiation of radicle
growth operates as a moisture-sensitive barrier to the
completion of germination. Thus, although the seed
is ‘ ready ’ to germinate, Ψ must exceed Ψb(G) before
radicle growth can be initiated. Such a mechanism
would avoid germination into seed-bed conditions
likely to be hostile to subsequent seedling growth. In
practice the interaction of this barrier and water
potential could determine the pattern of germination
curves under conditions where soil moisture varies
around Ψb(G). Indeed, the initiation to radicle
growth can be considered as a critical point at which
rain or irrigation is required to obtain uniform
germination in the field (Finch-Savage, 1990 a, b).
Only the end point of the germination process is
observed in experiments to determine germination
rate (reciprocal of time to germination) and therefore
the impact of a rate-limiting barrier cannot be
directly observed. However, the effect of such a
barrier has been shown in seed priming studies
where the progress of germination to radicle emerg-
ence is prevented (e.g. low water potential, Khan
(1992) ; abscisic acid, Finch-Savage & McQuistan
(1991) ; low temperature, Coolbear, Newell & Bryant
(1987)). Under these conditions germination is
prevented, but physiological changes occur which
result in more rapid germination when those barriers
are subsequently removed. This metabolic advance-
ment can be quantified using hydrothermal time
accumulated above values of Tb and Ψb(G) s
determined for the progress of metabolism which are
lower than those derived for radicle emergence in the
hydrothermal time model (Bradford & Haigh, 1994).
Thus the accumulation of this so-called ‘ hydro-
thermal priming time ’ is subsequently expressed as
more rapid germination (reduced hydrothermal
time) when the barrier to radicle emergence is
removed. Bradford (1995) speculates how the effect
of priming on germination rate might be predicted
by combining hydrothermal and hydrothermal
priming time.
The accuracy of prediction here using eqn (4) on
most occasions suggest that if seeds are sown into a
moist seed bed, which allows adequate imbibition,
they progress towards germination rapidly and are
relatively insensitive to the effect of Ψ in the seed bed
that remains above Ψb(G) until the initiation of
radicle growth when the condition Ψ " Ψb(G) must
be fulfilled. As the seed bed dries quickly at sowing
depth this will usually occur following rain or
irrigation. For example, at sowing 5, Ψ declined
during a dry spell following sowing into moist soil
and then, after some delay, germination occurred
rapidly when soil-water potential increased and Ψ "

514 W. E. Finch-Savage, J. R. A. Steckel and K. Phelps
100 (a)
80
Germination (%)
60
40
20
0·0
Water potential (MPa)
–0·5
–1·0
–1·5
0 100 200
Figure 7. Soil water potential and cumulative germination at sowings 2 (a) and 5 (b) in treatments receiving
pre-emergence irrigation. Predictions of germination time for each decile of the population according to eqns
1 (E) and 4 (*) subject to the conditions outlined in the text are also shown.
Ψb(G) (Fig. 7 b). Equation (4) described this pattern
satisfactorily. However, at sowing 2, Ψ fell below
Ψb(G) 2 d before germination occurred, it then
rained and soil-water potential increased just before
germination was first recorded. Under these cir-
cumstances, seeds surprisingly germinated according
to thermal time (Fig. 7 a), and germination rate was
apparently unaffected by Ψ ! Ψb(G). In eqn (4)
‘ time ’ only accumulates above Ψb(G), consequently
germination time was overestimated.
In general terms the above result suggests that the
Ψb(G) above which ‘ time ’ accrues in the period
leading up to the initiation of radicle growth is too
high. Arguably this period is more likely to be
controlled by the limits for metabolic advancement
rather than those determined for radicle emergence
itself. Therefore, lower Ψbs similar to those de-
termined for hydrothermal priming time (Bradford
& Haigh, 1994) might be more appropriate. In this
case, ‘ time ’ could accrue above a lower Ψb(G) for
metabolic advancement, although, the condition
Ψ " Ψb(G) for radicle growth initiation must still
be fulfilled before germination could occur. This
change would bring forward the predicted ger-
mination time at sowing 2 (Fig. 7 a). In most of the
other seed-bed environments where germination was
delayed and tended to occur only after rainfall, as
described for sowing 5 (Fig. 7 b), the prediction
would be little affected. At sowing 3 on unirrigated
plots (Fig. 4 g) neither hydrothermal time nor eqn (4)
predicted that germination was sufficiently far ad-
vanced to take advantage of the rainfall event which
actually brought about germination. These models
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(b)
300 0 100 200 300 400
Thermal time (°C d)
predicted that germination would occur after sub-
sequent rainfall events. The accumulation of ‘ time ’
above a lower Ψb(G) would have improved the
accuracy of this prediction.
The accuracy of germination predictions using
eqn (4) in most situations shows the importance of a
rate-limiting moisture sensitive step in predicting
germination in a variable seed bed. The comparative
lack of fit using hydrothermal time implies that time
accrues too slowly in the existing model under field
conditions. Results here suggest that the incor-
poration of metabolic advancement below Ψb(G)
into hydrothermal time theory, discussed by
Bradford (1995), is likely to considerably improve
predictions of germination time under variable field
conditions.
Sources of variation in seedling emergence
It is clear from the present work that when high
quality carrot seeds are sown under good hor-
ticultural practice any reduction in the number of
seedlings that emerge occurs after germination in the
soil. This is in agreement with studies on other crop
plants (Hegarty & Royle, 1978 ; Durrant, 1981).
When seedling emergence was greatly delayed
much of this delay occurred before germination.
This would be expected if the moisture-sensitive
rate-limiting step at the initiation of radicle emerg-
ence discussed above operates. If the seed bed is dry,
seeds would not germinate until sufficient rain fell to
increase soil Ψ above Ψb and the radicle would then
extend down into increasingly moist soil and thus, in

Carrot germination and post-germination growth
the absence of increased impedance, post-germi-
nation growth would not be delayed. Consequently
most delay in seedling emergence occurs in the
germination phase.
By contrast, results suggested that increases in the
spread of seedling emergence times seemed to be
largely manifested post-germination. This is con-
sistent with results that show carrot seeds selected as
fast germinating and grown without stress in lab-
oratory tests consistently grew faster and with greater
uniformity than slower germinating seeds in the
population (Finch-Savage & McQuistan, 1988).
Therefore, although not true for all species (Wheeler
& Ellis, 1991 ; Finch-Savage, 1995), clear differences
in intra-seed lot vigour can occur in this crop so that
slow germinating seeds might be inherently less able
to cope with post-germination seed-bed stresses. A
further complication is that, in the absence of light,
the rate of growth from smaller carrot seeds with less
reserves begins to slow down sooner and the final
seedling length they achieve is less (Tamet, Durr &
Boiffin, 1994).
In summary, the work presented suggests that
delays in seedling emergence occur largely in the
germination phase, which can be adequately
predicted using a modified threshold model, but
seedling losses and variation in the spread of seedling
emergence times within the population occur largely
during the post-germination growth phase.
              
We thank Mr C. I. McQuistan for technical help, Dr A.
Walker for assistance with his computer program and the
Ministry of Agriculture Fisheries and Food for funding
this work.

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