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The majority of experiments in plant biology use plants grown in some kind of container or pot. We conducted a meta-analysis on 65 studies that analysed the effect of pot size on growth and underlying variables. On average, a doubling of the pot size increased biomass production by 43%. Further analysis of pot size effects on the underlying components of growth suggests that reduced growth in smaller pots is caused mainly by a reduction in photosynthesis per unit leaf area, rather than by changes in leaf morphology or biomass allocation. The appropriate pot size will logically depend on the size of the plants growing in them. Based on various lines of evidence we suggest that an appropriate pot size is one in which the plant biomass does not exceed 1 g L-1. In current research practice similar to 65% of the experiments exceed that threshold. We suggest that researchers need to carefully consider the pot size in their experiments, as small pots may change experimental results and defy the purpose of the experiment.
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Pot size matters: a meta-analysis of the effects of rooting
volume on plant growth
Hendrik Poorter
, Jonas Bühler
, Dagmar van Dusschoten
, José Climent
and Johannes A. Postma
IBG-2 Plant Sciences, Forschungszentrum Jülich, D-52425, Germany.
INIA, Forest Research Centre, Department of Forest Ecology and Genetics, Avda A Coruña Km 7.5.,
28040 Madrid, Spain.
Corresponding author. Email:
Abstract. The majority of experiments in plant biology use plants grown in some kind of container or pot. We conducted a
meta-analysis on 65 studies that analysed the effect of pot size on growth and underlying variables. On average, a doubling of
the pot size increased biomass production by 43%. Further analysis of pot size effects on the underlying components of
growth suggests that reduced growth in smaller pots is caused mainly by a reduction in photosynthesis per unit leaf area,
rather than by changes in leaf morphology or biomass allocation. The appropriate pot size will logically depend on the size of
the plants growing in them. Based on various lines of evidence we suggest that an appropriate pot size is one in which the plant
biomass does not exceed 1 g L
. In current research practice ~65% of the experiments exceed that threshold. We suggest
that researchers need to carefully consider the pot size in their experiments, as small pots may change experimental results and
defy the purpose of the experiment.
Additional keywords: container volume, experimental setup, meta-analysis, pot size, plant growth, rooting volume.
Received 16 February 2012, accepted 11 May 2012, published online 15 June 2012
A large number of studies in plant biology focus on gene
expression, physiology or biomass production of individually-
grown plants. To this end, experiments are often conducted on
plants grown in some kind of container, from here on referred to as
pots. Pot size has received special attention in forestry (Carlson
and Endean 1976) and horticulture (Kharkina et al.1999), where
commercial companies can prot from choosing the smallest pot
volume that still delivers a product with an appropriate quality.
Occasionally, the issue of pot size has received attention in other
elds of plant biology. Arp (1991), for example, emphasised
that pot size might be an important issue in experiments that
considered the effect of elevated CO
on plants; and recently, a
discussion in the eld of ecophysiology has emerged, where
studies on the recognition of roots of neighbours are thought to
be confounded by pot size (Hess and De Kroon 2007).
Apart from the elds mentioned above, pot size seems to
have received little consideration in the scientic literature and
is regulary not reported in the materials and methods section of
publications. Nonetheless, it is an important issue. In most
laboratories there is large demand for growth chamber facilities
and the use of small pots generally implies more experiments or
increased replication. Space is probably less of an issue in most
glasshouses. However, the plant biology community currently
makes a great effort to develop automated systems for plant
phenotyping (Granier et al.2006; Nagel et al.2012). These
high-throughput systems allow for the handling of many plants,
which automatically implies that spatial demands on growth
facilities will increase. The use of small pots has the additional
advantage that it does not exceed the capabilities of the transport
robots to move the weight of pot plus plant.
The use of small pots for research purposes may also have
disadvantages, which are more related to biological constraints.
A small container implies a small quantity of soil or other
substrate and thereby, almost invariably, a reduction in the
availability of water and nutrients to the plant. In addition to
reduced resource availability, pots generally impede root growth.
Many species easily produce roots of more than 1 m in length
(Jackson et al.1996), even at a relative young age (Drew 1975;
Fusseder 1987), thereby exceeding the dimensions of most
containers used. Large plants in small pots may have a large
fraction of roots pot-bound, with all kind of secondary
consequences (Herold and McNeil 1979). In this paper we
explore the consequences of the choice of pot size for plants
studied for experimental purposes. We rst analysed in a
quantitative way to what extent pot size affects plant growth in
the studies that explicitly considered this factor. We did so by a
meta-analysis of 65 studies reported in the literature. Second,
we reported on some of the morphological and physiological
components that might explain the observed growth patterns.
Functional Plant Biology,2012, 39, 839850 Review
Journal compilation CSIRO 2012
Third, we analysed whether there is a threshold in the plant
size : pot size relationship above which growth is affected and
compared the data for the current pot-size experiments with a
more general database on plant research and to plants grown in
elds. Based on these comparisons we suggest what might be an
appropriate pot size for a given experiment.
1. Effect of pot size on plant biomass
To analyse the effect of rooting volume on plant growth, we
screened the literature of the last 100 years and arrived at a total
of 65 publications where this factor was studied in pots and
~10 where rooting volume was constrained in hydroponically-
grown plants (see Appendix 1). For this analysis we will focus
mainly on the results for the pot-grown plants. A description of
the methodological approach is given in Appendix 2. The effect
of pot size has been studied in a wide range of pot volumes, with
values ranging from 5 mL for a herbaceous greenhouse crop
(Bar-Tal and Pressman 1996), to 1700 L for trees growing over a
period of several years (Hsu et al.1996). The range of pot sizes
used within the various experiments was also large, varying by a
factor 2 at least and a factor 35 at most. A clear example in our
compilation is the experiment by Endean and Carlson
(1975), who followed the growth of Pinus contorta Douglas
over time. In the very beginning of the experiment, plants grew
presumably well in all pot sizes, but after 4 weeks of growth
biomass was already reduced in the smallest pot volume
(Fig. 1a), and by the last harvest, none of the pots seemed
large enough to ensure unrestricted growth.
In the full compiled data set the picture is similar: in almost
all of the experiments considered plants increase in weight with
every larger pot that is used (Fig. 1b). Most experiments
fall between the dotted lines, which indicate a xed ratio of
biomass to pot volume of 2 and 100 g L
, respectively, for the
lower and higher line. Only a few experiments with trees in large
containers exceed the 100 g L
. As all experiments start with
small seedlings, plants move over time from the bottom part of
the graph upwards (cf. Fig. 1a). Clearly, the experiments at the
upper right end of Fig. 1bare also the ones that were planned
to last for the longest time, 2 years in case of the experiment with
the largest pots (Hsu et al.1996), 5 years in case of the experiment
with the highest ratio (Bar-Yosef et al. 1988).
Theoretically, doseresponse curves should level off at
higher pot sizes, as in Fig. 1a. However, this is not the case
for many of the studies plotted in Fig. 1b, which mostly show
linear responses. We discuss this further in section 4. To scale all
data to an equal change in pot size, we calculated for each
species in each of the experiments the average slope of the
log-transformed mass versus pot size relationship and derived
from those values an easy to understand expression that indicates
the percentage increase in biomass for a 100% increase in pot
size (Appendix 3). On average, plants increased 43% in mass for
every doubling in pot size (Fig. 2,P<0.001), with no signicant
differences in response between herbaceous and woody species.
These response values are substantial and imply that plants
are likely to be over 3 times larger in 2 L pots than in pots of
0.2 L. Given these differences, it is clear that pot size should
be given careful consideration during the planning phase of
an experiment.
2. Effect on components of the carbon-budget
What is the cause of the growth retardation in smaller pot sizes?
As clearly shown by Endean and Carlson (1975), the effect of pot
size on biomass gradually increases throughout (the later part of)
the experimental period. The rate by which biomass of individual
plants is accumulated is proportional to the size of the plant and
conveniently described by the relative growth rate (RGR, the
rate of increase in biomass per unit biomass present; Evans
1972). The differences in RGR of plants growing in different
pot sizes are always smaller than the differences in biomass at the
end of the experiment. This implies that the physiological and
morphological factors that underlie the variation in biomass will
also be affected to a smaller extent than biomass itself (Poorter
et al.2012a). We analysed growth in terms of the plants carbon
economy, using a top-down approach where the RGR is
factorised into three components:
0.01 0.10 1.00
0.01 0.10 1.00 10.00 100.00 1000.00
10 000
100 000
Total dry mass (g)
Pot size (L)
Fig. 1. (a) Doseresponse curves of total plant dry mass of Pinus contorta
as dependent on pot size. Plants were grown in pots with six different sizes
and harvested at several times during a 20-week period. Numbers indicate
the time of harvest after sowing, in weeks. Data from Endean and Carlson
(1975), except for seed mass (week 0), which was taken from McGinley et al.
(1990). (b) Summary of total biomass data of plants grown in pots of
various sizes, as reported in the literature (65 experiments on 69 species,
see Appendix 1). Each line connects the observations of one species or
genotype in one experiment. Values in red indicate woody species, in blue
herbaceous species. Dotted lines indicate a total plant biomass per unit pot
volume of 2 (lower line) and 100 (upper one) g L
. Additional, unpublished
data were obtained for Barrett and Gifford (1995) and Liu and Latimer
(1995), as well as from L. Mommer and H. de Kroon, and R. Pierik.
840 Functional Plant Biology H. Poorter et al.
according to Evans (1972). Here, SLA denotes specic leaf area
(leaf area per unit leaf dry mass; m
), LMF the relative
allocation of biomass to leaves (leaf mass fraction, g leaf g
and ULR is a parameter that indicates the growth rate per unit
leaf area (unit leaf rate, g m
). ULR is basically the net
result of carbon gain through photosynthesis, corrected for the
rate of respiration in the whole plant and the C-content of the
newly added biomass. ULR and photosynthesis per unit leaf
area are often strongly positively correlated (Poorter 2002).
A characteristic of RGR is that an absolute difference in RGR
between treatments causes a relative difference in biomass over
time (Poorter and Navas 2003). However, in the case where we
want to understand the reason for the difference in growth and
only have fragmented information, it is more amenable to analyse
the proportional differences in RGR relative to that of the three
growth parameters that underlie RGR (Eqn 1). Only a few
experiments report data on these underlying components, but
the information we could gather points into the following
direction (see Fig. 2and Fig. S1, available as Supplementary
Material to this paper): a doubling in pot size increases RGR by
~5%. Consequently, each of the growth parameters at the right-
hand side of Eqn 1 may change by a very small proportion, or
one of them by a somewhat larger proportion in the range of
5%. At the nal harvest, SLA increased somewhat in some
experiments and decreased in others. We could quantitatively
compare the various experiments by calculating the percentage
increase in SLA with a 100% increase in pot size and found that
taken over all experiments, this variable did not deviate
signicantly from zero (Fig. 2). Most, but not all experiments
with root restriction in hydroponics conrm this response (e.g.
Carmi et al.1983; Kharkina et al.1999; but see Tschaplinski and
Blake 1985).
Allocation patterns are more frequently reported, generally
as the biomass ratio between shoot and root. We prefer a
classication in at least three plant organs (Poorter et al.
2012b) and therefore use LMF, SMF and RMF, which are the
fractions of total vegetative biomass invested to leaves, stems
and roots respectively. The relatively scarce information indicates
that LMF is not affected (Fig. 2a), whereas SMF increased
slightly but non-signicantly (0.05 <P<0.10) with pot size.
More information is present on RMF (or the root : shoot ratio).
Reviews are not equivocal in their evaluation. NeSmith and
Duval (1998) conclude that RMF often does not change with
pot size. In contrast, Hess and De Kroon (2007) expect RMF
to increase in larger pots. We found that over ~80 species
experiment combinations, RMF decreased on average by a small
but signicant extent (4% with a doubling in pot size, P<0.05)
We nd no clear differences between woody and herbaceous
plants in this respect (data not shown).
With two of the variables of Eqn 1 unaffected, we might
expect that the observed differences in growth rates between
plants grown in different pot sizes are caused largely by a
change in net photosynthesis and hence we would expect a
stimulation of ~4% in this variable. Some experiments report
a more than 30% higher rate of photosynthesis with a 100%
increase in pot size (Robbins and Pharr 1988; Ronchi et al.
2006; Fig. S1g). This is far more than the expected 4%. As pot
size stress builds up gradually over time, results may depend on
the actual timing of the measurements during the experiment.
Moreover, obtained rates will also depend on the specic leaf
measured and the time of the day that the measurement was
taken. The growth parameter ULR is generally closely linked to
the rate of photosynthesis and provides an estimate that is
integrated over the whole plant and the whole growth period
under consideration (Poorter 2002). A 2-fold difference is not
reported in the literature (Fig. S1f), but the pot-size effects on
ULR are signicantly greater than zero and larger than for SLA
and LMF (Fig. 2). The median increase in photosynthesis and
ULR with a doubling in pot size is quite comparable to the
median increase in RGR. Hence, we conclude that with the
% Change with 100%
increase in pot size
TDM (g) or RMF (g g–1)
in large/in small volume
Fig. 2. (a) Distribution of the percentage increase in a range of growth-
related traits when pot size doubles. The data are from a range of
experiments described in literature (Appendix 1). The distribution is
characterised by box and whisker plots, where the boxes show the 25th
and 75th percentile and the whiskers the 10th and the 90th percentile. The
median is represented by the line in the box and given as a number above the
box plot. The total (rounded) number of species experiment combinations
on which the boxplots are based are TDM (total dry mass): 90; RGR (relative
growth rate): 15; SLA (specic leaf area): 20; LMF (leaf mass fraction):
35; SMF (stem mass fraction): 30; RMF (root mass fraction): 80; ULR
(unit leaf rate): 5; PSa (rate of photosynthesis per unit leaf area): 15; LNC
(leaf nitrogen concentration): 15. The signicance level of a test whether
the observed distribution deviates signicantly from 0 is given above the
respective box plots: ns, not signicant;
, 0.05 <P<0.10; *, P<0.05;
**, P<0.01; ***, P<0.001. (b) Distribution of ratios in TDM and
RMF, taken from pot size studies (values for the largest pot relative to
those for the smallest pot within each experiment, n= 65) and studies
where rooting volume was constrained for hydroponically-grown plants
(ratio unconstrained/constrained, n= 11). The results of t-tests between the
two groups are shown between the boxplots.
Pot size matters Functional Plant Biology 841
limited and fragmentary evidence yet available, net
photosynthesis is likely to be the process that is strongest
affected by pot size and may explain best the observed pot
size effect on biomass (Fig. 1b). Additional support comes
from experiments where photosynthesis recovered quickly
after plants were repotted in larger rooting volumes (Herold
and McNeil 1979).
3. What mechanism could explain a reduced
photosynthesis in smaller pots?
Several factors could explain the reduced rate of photosynthesis
and thereby growth in smaller pots. A rst possible explanation
is that containers of smaller dimensions can be placed at a higher
density, with less light available for each shoot and hence, a
lower rate of photosynthesis. Although this could be the case in
some of the compiled experiments, strong pot size effects on
biomass and photosynthesis are generally also observed when
density is specically controlled for (e.g. Endean and Carlson
1975; Robbins and Pharr 1988; NeSmith et al.1992; Climent
et al.2011).
In the type of experiments that we included in our meta-
analysis, a smaller pot size will inadvertently decrease the total
nutrient content in the pot. Low nitrogen and phosphorus
availability are known to decrease photosynthesis (e.g.,
Sinclair and Horie 1989; Lynch et al.1991) and growth and
increase the root mass fraction (Poorter et al.2012b). Thus,
lower resource supply could form a plausible explanation. We
calculated the response of leaf nitrogen concentrations to changes
in pot size, expecting to see an increase if nutrient availability
would explain the pot size effect. On average, there was a slight,
but non-signicant increase in leaf nitrogen concentrations with
pot size (Fig. 2), suggesting that this factor cannot completely
explain the observed differences in photosynthesis or growth.
Similar results were found for phosphorus (Krizek et al.1985).
This conclusion is to a certain extent supported by observations
on hydroponically-grown plants, which have decreased
photosynthesis and growth (Fig. 2b) when the root volume
was restricted, despite a continuous high supply of nutrients.
However, unlike plants grown in pots, hydroponically-grown
plants do not show an increased RMF when restricted (Fig. 2b).
As increased RMF is a good indicator for nutrient stress, we
presume that nutrient limitation in small pots is still a factor,
although we cannot exclude possible allometric effects which
could explain a larger RMF in smaller plants as well (Poorter et al.
Water is the other commodity that may be in short supply.
Small pots could negatively impact the water status of plants
as they have a reduced total water holding capacity and
will, therefore, dry out more quickly (Tschaplinski and Blake
1985) and at severe stress levels increase RMF (Poorter et al.
2012b). Ray and Sinclair (1998) demonstrated with their
drought experiment that soil in small pots dries out faster and
thereby caused more severe drought stress in plants. However,
pot size does not necessarily affect stomatal conductance or
leaf water potential (Ronchi et al.2006) and as for
nutrients with plants growing in hydroponics there is still a
clear effect of root connement, even though water availability is
not restricted.
Besides resource availability, the temperature of the
rooting volume could be affected by pot size (de Vries 1980).
Pots can intercept a substantial amount of solar radiation
especially in experimental gardens and glasshouses, which
may increase the soil temperature at the edge and eventually
in the middle of the pot if no precautions are made (Martini
et al.1991;Xuet al.2001). Small pots have greater surface
areas relative to their volume and thereby heat up more
quickly. Townend and Dickinson (1995) measured 5C
higher day temperatures in 0.19 L pots compared with 1.9 L
pots. Keever et al.(1986) suggest that the greater temperature
uctuations in small pots may explain the reduced growth of
the plants. High temperatures in the pot may have several direct
(respiration, root growth) and indirect (through increased
microbial activity) effects on plant growth. Pot temperatures
are rarely reported so it is difcult to evaluate how often
temperature differences between pots contribute to reduced
growth. However, given that growth reductions also have been
observed in hydroponically-grown plants suggests that
temperature differences alone cannot explain the observed pot
effects either.
If neither nutrient or water availability nor higher temperatures
can (fully) explain the pot size effects on photosynthesis and
growth, it could be that root connement per se may cause growth
retardation, with reduced photosynthesis as a consequence.
Root growth is known to respond directly to impedance.
Impeded roots stay shorter whereas the initiation and growth
of side branches increases (Bengough and Mullins 1991; Falik
et al.2005). Furthermore, Young et al.(1997) showed that within
10 min of increasing the impedance to root growth, leaf expansion
rate is reduced. This suggests that some kind of signal may
regulate shoot growth when a large proportion of the roots are
impeded. The actual signal for such a response remains as yet
unknown. Possibly a reduced sink strength of the root system
could cause a direct negative feedback on photosynthesis (Paul
and Pellny 2003). Alternatively, a specic rootshoot signal is
involved (Jackson 1993).
A crucial point in the evaluation of the lastly discussed
option is knowledge on the actual distribution of roots within
the pots. Although the vertical root distribution is relatively
easily measured (Price et al.2002; Suriyagoda et al.2010),
analysis of the horizontal distributions is more complicated.
Using non-destructive magnetic resonance imaging (MRI), we
followed the root development of Hordeum vulgare L. and
Beta vulgaris L. plants over time in three dimensional space.
Representative nuclear magnetic resonance (NMR) images of
root systems at the end of the experiment are shown in Fig. 3.
We calculated the percentage of roots that was located in
the inner half of the soil volume, furthest away from wall and
bottom and the percentage of roots present in the outer 4 mm of
the pot. Only 2025% of the root biomass was in the inner part
of the pot (Table 1), whereas ~50% was found in the outer 4 mm
(20% of the total volume). The proportional distribution
remained remarkably constant over time. Hence, if these
observations have wider validity, we conclude that a relatively
large fraction of roots is close to the edge of the pot, where
unfavourable environmental conditions, for example, large
temperature uctuations and impedance of the pot wall may
negatively impact growth.
842 Functional Plant Biology H. Poorter et al.
4. When does pot size limitation starts?
In sections 1 and 2 we considered for each experiment the
overall effect on plant growth, morphology and physiology
when pot size was doubled. However, it is to be expected that
a plant of a given size will be constrained more in a small than in a
large pot. That is, young plants are initially not affected by pot
size, but as plants grow older, the pot size effect becomes more
pronounced, even in medium-sized pot volumes (Fig. 1a). When
experiments last for sufciently long time, even the largest pot
size might not be large enough for unrestricted growth, i.e. the
saturating part of the curve extends beyond the largest pots used.
For the experimental data this implies that the relationship
becomes close to linear again. In fact, many of the curves in
Fig. 1bshow a linear relationship. For experiments where only
two pot sizes were used, it is impossible to deduce whether the
response of the plants is indeed linear or not. But even in many of
the other experiments in Fig. 1bno clear saturation is shown.
What is the reason for that?
One objective way to relate plant and pot size across
experiments is to calculate the plant biomass that is present at
a given volume of rooting space. This variable, for which we use
BVR as an acronym (total plant biomass : rooting volume ratio;
), has, to our knowledge, been used only by Kerstiens and
Hawes (1994). BVR values vary widely and ranged in our
database from as low as 0.01 in work by Climent et al.(2008)
to over 300 g L
in work reported by Biran and Eliassaf (1980).
The median value in the pot size experiments is around 9.5 for
experiments both with herbaceous and woody species (Fig. 4b).
We tested whether the BVR could explain the form of the
doseresponse curves in the 65 experiments shown in Fig. 1b,
by calculating for each point what the slope of the doseresponse
curves was, as well as the BVR. In order to be consistent with
Fig. 2, we derived the percentage increase in biomass with pot
size doubling for these data as well. We found that very few
experiments had BVR values lower than 2 g L
(Fig. 4b). Hence,
for this part of the analysis we included not only data of the last
harvest, but also data of earlier harvests where available. This may
imply that not all data points in the analysis are formally
independent, but it increases the power of our analysis in this
crucial range. We binned all values in ve BVR ranges and show
the resulting distribution in Fig. 5a. Estimation of slopes is
always more challenging than determining the absolute values
per point and this may be one of the reasons that there is
considerable variation within each category. In the category
with a BVR between 1 and 2 g L
the effect of pot size is
clearly noticeable. Pot size effects are saturating when BVR
values exceed the 2 g L
Another way to obtain a greater insight into the relationship
between plant biomass and pot size is to express the biomass of
Fig. 3. (a) NMR image of a Hordeum vulgare plant grown in a pot with a
volume of 1.3 L for 44 days. Roots in the inner 50% of the soil volume
(furthest away from wall and bottom) are colour-coded yellow, roots in the
outer 50% blue. The stem part that was masked from the analysis is shown in
red. (b) Idem for a Beta vulgaris plant 48 days after sowing. The developing
storage root is colour-coded red and was not included in this case.
Table 1. The proportion of the root mass that is present in the inner 50% of the pot volume (more than 12.5 mm from
the wall or bottom) and in the space less than 4 mm from wall or bottom of the pot, for Hordeum spontaneum and
Beta vulgaris plants growing in 1.3 L pots
Values are based on six plants followed over time. Time is days after sowing. Standard error of the mean was on average 1.6
and 2.9 percentage points for the inner half and outer 4 mm respectively
Hordeum vulgare Beta vulgaris
Root mass in
inner half (%)
Root mass in
outer 4 mm (%)
Root mass in
inner half (%)
Root mass in
outer 4 mm (%)
26 32.0 43.2 31 20.3 53.5
28 28.9 50.0 33 20.8 53.1
31 17.7 55.3 34 21.5 54.1
33 21.3 52.7 39 22.9 52.6
34 23.1 51.7 41 23.0 52.3
39 25.2 48.7 48 22.3 52.0
41 28.2 48.0 –– –
44 32.4 44.7 –– –
Pot size matters Functional Plant Biology 843
plants grown at various pot sizes relative to the biomass at
the largest pot size and plot these values against the BVR
(Fig. 5b). Experiments where pot sizes are limiting growth
throughout the full range of pot sizes are characterised by lines
that decline linearly with BVR. However, as long as biomass is
not affected, the line will remain around 1 and only drop at greater
BVR values when pot size starts to reduce growth. For the few
experiments where this was the case, we could show that this
inection point occurred somewhere between 0.2 and 2 g L
(Fig. 5b). Thus, from both the full sets of experiments compiled,
as for the more detailed analyses over time, we derive that pot
size effects are particularly strong when BVR values are greater
than 2 g L
5. How do these BVR values relate to other
experiments and the eld?
As mentioned in section 1, the range of pot sizes used in this
compilation is large. The median value is around 0.9 L (Fig. 4a).
How does that compare to common practice in ecophysiological
experiments? This will partly depend on the species studied.
Arabidopsis, for example, is generally grown in much smaller
pots (with an interquartile range of 0.080.21 L) than Zea
mays L. (1.85.0 L). An overall impression of used pot sizes
can be obtained from the metaphenomics database described
by Poorter et al.(2010), where the response of ~900 different
species to 12 different environmental factors is compiled for a
total of ~800 experiments from the literature. The median pot
size used in that compilation of experiments turns out to be
~23 times larger than those in the pot size studies (Fig. 4a),
whereas the median BVR value is ~4-fold lower (Fig. 4b). Hence,
we conclude that most studies on pot sizes have focussed on
relatively small pots and have grown plants to larger sizes
than is common in ecophysiological experiments. In contrast,
experiments in plant biology generally use relatively larger pots
and harvest plants at younger stages, when the BVR value is still
below 8 g L
Pot size (L)
BVR (g L
Pot size
Herb. Herb.
Fig. 4. (a) Distribution of pot volumes as represented in the current meta-
analysis of pot size studies and in a compilation of ~800 studies on the effect
of 12 environmental factors on growth and related ecophysiological traits
(meta-phenomics database, Poorter et al.2010). (b) Total plant biomass : pot
volume ratios (BVR) in the current meta-analysis and the meta-phenomics
database. The distribution is characterised by boxplots (see legend Fig. 2).
Blue boxes indicate the values for herbaceous species, red ones for woody
species. Numbers above/below each box show the median values.
0.1 1.0 10.0
Scaled total dry mass (g g–1)
>201–2 2–5 5–20<1
% Change in biomass with 100%
increase in pot size
Fig. 5. (a) Distribution of the percentage change in biomass with a
doubling in pot size for a total of ~80 experiment species combinations.
Distributions are characterised by boxplots and given separately here for
ve classes of BVR (total plant biomass : pot volume ratio). The ANOVA was
highly signicant (P<0.001), with an r
of 0.22. The (rounded) number
of observations in the ve classes are 30, 30, 90, 200 and 80 respectively.
(b) The total dry mass of plants in various pot sizes, scaled to the biomass
of the plants in the largest pot, plotted against BVR. Different lines indicate
data from different experiments or different harvests within an experiment.
844 Functional Plant Biology H. Poorter et al.
Most experiments with pots are conducted to eventually
understand how (agro-)ecosystems function. It may therefore
be relevant to consider what normal BVRvalues are in the eld.
Maximum dry matter yield of major crops varies between
10003000 g m
(Unkovich et al.2010; assuming 20%
biomass in roots). If we assume a rooting depth of 1 m, this
would correspond to BVR values in the range of 13gL
. Given
that at least half of the DM production takes place during
grain lling, we can expect that during the vegetative state, the
BVR value will not exceed 1.5 g L
. Similar calculations for
natural ecosystems are more difcult as large variation exists
in root depth and the standing biomass. Given a rooting depth
of 0.35 m (Nagel et al.2012) and a density of 500 plants per m
aeld of Arabidopsis thaliana Heynh. plants of 0.1 g dry mass,
would have a BVR of 0.15 g L
. Although we realise that plants
in the eld experience conditions that are very different from
those where plants are grown singly in pots in controlled
conditions, we conclude from these rough estimates that BVR
values around 1 are of the same order of magnitude as those of
vegetative plants in the eld.
6. Does pot size affect experimental conclusions?
Up to now we have considered the effect of pot size per se. Most
researchers are also interested in whether the outcome of their
experiments is affected by the choice of the pot size. Arp (1991)
was one of the rst to draw attention to the fact that pot size
might restrict the response to elevated CO
. This would limit the
possibilities to draw conclusions from experiments that have been
conducted in this eld.
The analysis by Arp (1991) was a compilation of different
studies that worked with different pot sizes. Kerstiens and Hawes
(1994), however, published a meta-analysis of the results of a
range of studies with trees in which they show that the biomass
responses to elevated CO
did not correlate with pot size, or even
decreased with a BVR over 18 g L
. This suggests that small
pot sizes do not reduce responses to elevated CO
However, in both meta-analyses the evidence could only be
circumstantial, as pot size was not an experimental factor in
the compilations. Here we analysed experiments where pot size
was specically included in the experimental design, not only
for interaction with CO
but also for nutrients, water and
irradiance (Fig. 6). Given that nutrient and water availability
already increase when pot size is increased, we would expect
less additional effect on plant growth if more nutrients or water
were supplied. However, we would expect increasingly stronger
growth responses with larger pot size when light or CO
be increased, simply because of the higher demand for nutrients
and water in larger plants. Although some of the experiments do
indeed follow the expected trend, results are not equivocal. As
results depend on the variability in at least four different harvests,
the number of experiments is likely too small to draw any strong
Besides possible interactions with abiotic factors, interactions
with biotic factors have been shown. For example, vesicular
arbuscular mycorhizae (VAM) infection rates, which would
normally increase with reduced nutrient availability, are
reduced in small pots (Bååth and Hayman 1984; Koide 1991).
As a consequence, VAM colonisation is less benecial for
nutrient uptake and results in smaller growth increases when
small pots are used (Kucey and Janzen 1987; Koide 1991).
Similarly, Baldwin (1988) showed that pot-bound Nicotiana
plants do not respond to leaf damage, whereas repotted plants
do. Thus, although we cannot draw a rm conclusion here we
suggest that the use of small pots with crowded roots carries a
substantial risk of inuencing the experimental results.
7. Other considerations
This analysis focussed on the effect of pot volume on plant
growth. However, choosing a pot for an experiment not only
includes choosing the right volume, but also the right shape.
Although shape is less important than volume (McConnaughay
et al.1993), shallow and deep-rooting species may respond
differently to the actual diameter and height of the pots, at
equal pot volume (von Felten and Schmid 2008). Pot height is
also an important factor in determining the free-draining water
content of pots and thereby the water potential as well as the
oxygen availability in the pots (Passioura 2006).
An alternative to standard plastic pots are containers that have
ribbed inner sides and small air holes. Such containers promote
self-pruning of roots close to the holes, which avoids root
spiralling and promotes development of lateral roots (Rune
2003). Not only shape, but also the material (Bunt and
Kulwiec 1970) and the colour of the pot (Markham et al.
2011) may affect plant growth, mainly through their effect on
soil and root temperature. For a broader discussion on the use of
pots for growing plants in the context of experimental setup see
Poorter et al.(2012a).
A meta-analysis of the effects of pot size on growth shows
that on average a doubling of the pot size results in 43% more
biomass. In most cases reduced growth in small pots will be
caused by a reduction in net photosynthesis. It is the plant
0.1 1 10 100
Ratio biomass high vs. low (g g–1)
Pot size (L)
Fig. 6. Interaction of pot size with various abiotic factors. Results are
given for total plant biomass and are expressed as the ratio between the
biomass at a high level of that factor and the lower level. Data are from
Bilderback (1985), Thomas and Strain (1991), McConnaughay et al.(1993),
NeSmith (1993), Ismail et al.(1994), Nobel et al.(1994), Barrett and Gifford
(1995), Will and Teskey (1997), Houle and Babeux (1998), Centritto (2000)
and R. Pierik (unpublished data).
Pot size matters Functional Plant Biology 845
mass per unit rooting volume that is relevant rather than pot size
per se. Large plant mass per pot volume not only reduces growth
of plants but also carries the risk of inuencing the
relative differences between treatments. We conclude that it is
important for researchers to minimise such effects by choosing
pots that are large enough for their plants, even at later stages of
growth. Our advice is to avoid plant biomass to pot volume
ratios larger than 2 gL
and preferably work with plant and pot
sizes where this ratio is <1.
Damian Barrett, Phil Grime, Joyce Latimer, Liesje Mommer and Ronald
Pierik kindly provided (additional) unpublished data for this analysis.
Christian Heinemann enlightened us with the mathematical aspects and
comments of Liesje Mommer, Thijs Pons, Marc Faget as well as two
anonymous reviewers on a previous version of the ms are greatly
appreciated as well.
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Appendix 1. List of references used for the meta-analysis
A. Pot studies
Cris and Stout (1929) Agric. Exp. Stat. MSC, No 6.; Stevenson (1967) Can. J. Soil Sci. 47, 163174; Endean and Carlson (1975) Can.
J. For. Res. 5,5560; Hocking and Mitchell (1975) Can. J. For. Res. 5, 440451; Carlson and Endean (1976) Can. J. For. Res. 6,
221224; Herold and McNeil (1979) J. Exp. Bot. 30, 11871194; Biran and Eliassaf (1980) Sci. Hortic. 12, 385394; Peterson et al.
(1984) Agron. J. 76, 861863; Bilderback (1985) J. Env. Hortic. 3, 132135; Krizek et al. (1985) J. Exp. Bot. 36,2538; Carmi
(1986) Field Crops Res. 13,2532; Hanson et al. (1987) HortSci. 22, 12931295; Kucey and Janzen (1987) Plant Soil 104,7178;
Ruff et al. (1987) J. Amer. Soc. Hort. Sci. 112, 763769; Tilt et al. (1987) J. Amer. Soc. Hort. Sci. 112, 981984; Bar-Yosef et al.
(1988) Plant and Soil 107,4956; Robbins and Pharr (1988) Plant Physiol. 87, 409413; Bar-Tal et al. (1990) Agron. J. 82, 989995;
Gurevitch et al. (1990) J. Ecol. 78, 727744; Koide (1991) Oecologia 85, 389395; Latimer (1991) HortScience 26, 124126;
Martini et al. (1991) J. Am. Soc. Hort. Sci. 116, 439445; Simpson (1991) North. J. Appl. For. 8, 160165; Thomas and Strain (1991)
Plant Physiol. 96, 627634; Dubik et al. (1992) J. Plant Nutr. 15, 469486; NeSmith et al. (1992) J. Plant Nutr. 15, 27632776;
Samuelson and Seiler (1992) Env. Exp. Bot. 32, 351356; Beeson (1993) J. Amer. Soc. Hort. Sci. 118, 752756; McConnaughay
et al. (1993) Oecologia 94, 550557; NeSmith (1993) J. Plant Nutr. 16, 765780; Ismail et al. (1994) Aust. J. Plant Physiol. 21,
2335; Menzel et al. (1994) J. Hortic. Sci. 69, 553564; Nobel et al. (1994) Physiol. Plant. 90, 173180; Ran et al. (1994) Agron. J.
86, 530534; Barrett and Gifford (1995) Aust. J. Plant Physiol. 22, 955963; Liu and Latimer (1995) HortSci. 30, 242246; Mandre
et al. (1995) J. Amer. Soc. Hort. Sci. 120, 228234; Agyeman et al. (1996) Ghana J. For. 2,1424; Hsu et al. (1996) HortSci.31,
11391142; Huang et al. (1996) Plant and Soil 178, 205208; Ismail and Noor (1996) Sci. Hortic. 66,5158; McConnaughay et al.
(1996) Ecol. Appl. 6, 619627; Giannina et al. (1997) Acta Hortic. 463, 135140; Van Iersel (1997) HortSci. 32, 11861192; Will
and Teskey (1997) Tree Physiol. 17, 655661; Houle and Babeux (1998) Can. J. Bot. 76, 16871692; Nishizawa and Saito (1998)
J. Amer. Soc. Hort. Sci. 123, 581585; Ray and Sinclair (1998) J. Exp. Bot. 49, 13811386; Boland et al. (2000) J. Amer. Soc. Hort.
Sci. 125, 135142; Centritto (2000) Plant Biosystem.134,3137; Haver and Shuch (2001) Plant Growth Reg. 35, 187196; Yeh and
Chiang (2001) Sci. Hortic. 91, 123132; Aphalo and Rikala (2003) New Forests 25,93108; Loh et al. (2003) Urban For. Urban
Green 2,5362; Ronchi et al. (2006) Funct. Plant Biol. 33, 10131023; Arizaleta and Pire (2008) Agrociencia 42,4754; Chirino
et al. (2008) For. Ecol. Man. 256, 779785; Climent et al. (2008) Silvae genetica 57, 187193; Goreta et al. (2008) HortTechnology
18, 122129; Kurunc and Unlakara (2009) New Zeal. J. Crop and Hortic. Sci. 37, 201210; Oztekin et al. (2009) J. Food Agric. Env.
7, 364368; Climent et al. (2011) Eur. J. Forest Res. 130, 841850; Nord et al. (2011) Funct. Plant Biol. 38, 941952; Mommer and
De Kroon (pers. comm.); Pierik (pers. comm.).
B. Hydroponically grown plants
Richards and Rowe (1977) Ann. Bot. 41, 729740; Carmi et al. (1983) Photosynthetica 17, 240245; Tschaplinski and Blake (1985)
Physiol. Plant. 64, 167176; Hameed et al. (1987) Ann. Bot. 59, 685692; Peterson et al. (1991) J. Exp. Bot. 42, 12331240; Thomas
(1993) Plant Growth Reg. 13,95101; Ternesi et al. (1994) Plant Soil 166,3136; Bar-Tal et al. (1995) Sci. Hortic. 63, 195208; Bar
and Pressman (1996) J. Amer. Soc. Hort. Sci. 121, 649655; Kharkina et al. (1999) Physiol. Plant. 105, 434441; Xu et al. (2001)
J. Plant Nutr. 24, 479501.
848 Functional Plant Biology H. Poorter et al.
Appendix 2
For this meta-analysis we screened the literature of the last 100 years. A total of 63 publications plus two additional unpublished
experiments dealt with plants grown in pots of various sizes, 11 with plants grown in hydroponics with different levels of root
connement. The references are listed in Appendix 1. As we were interested in not only the physical aspect of container volume, but
also in the resources that come with it, we compared pot size treatments based on size, including the possible additional benets of
increased nutrient and water availability. The experiments with hydroponically-grown plants are not included in the main analysis.
Differences in the shape of the pots were not independently analysed either. In several publications only pot diameter was reported.
For a sample of 30 different pots ranging in diameter (Ø) from 7 to 40 cm we derived an estimate for pot volume (V) based on the
empirical equation V=p(Ø/2)
(0.46 + 0.8397 Ø0.002307 Ø
). If additional data were missing, we assumed pots to be lled
with substrate up to the rim.
For each species or genotype in a given experiment, we determined the biomass at the last harvest and separated this variable in
biomass of leaves, stems, roots and reproductive mass as far as data were provided. To capture the importance of pot size for growth we
calculated for each experiment the proportional increase in total plant dry mass relative to the proportional increase in pot size, by
calculating the slope of the lines that were tted through the observed log-transformed plant masses and pot volumes, separately for
each species in each experiment. Because experiments differed in the range of pot size used, we scaled these slopes in such a way that
the number reects the percent change in biomass (or another variable) given a doubling in pot size (see Appendix 3 for more details).
For a more detailed analysis, log-transformed data from experiments which included three or more pot sizes were also tted with a
saturating equation, as described in Appendix 3 and calculated with the nls procedure in R (R Development Core Team 2011).
Appendix 3
Let V
and V
be pot volume 1 and 2 and B
and B
the total plant dry biomass that is observed at the respective pot volumes:
because proportional differences are the focus of interest, we calculate the slope of the line that connects these points as:
Suppose we want to know the fraction fby which plant mass increases when pot size doubles. Then
Assuming a common slope sover the whole trajectory of pot masses considered, this results then in
and, after rearrangement
The same procedure applies if slope s is determined by linear regression through more than two points.
To relate the slope of the line to the observed values of total plant mass per unit pot volume (BVR), we took the following approach:
In cases where an experiment consisted of two pot sizes, a linear regression as above was calculated and the resulting slope attributed
to both points. In case an experiment consisted of more than two pot sizes and a second order polynomial showed no signicant
saturating trend, we tted a straight line through all points. Otherwise, a saturating curve was tted through the points, of the form
with the constraint that a,band cshould be positive. The slope of the line at each pot size is then given by the derivative:
S¼ac b
After the slope and fwere calculated for each pot volume in each experiment, data were binned in ve categories of BVR.
Pot size matters Functional Plant Biology 849
Appendix 4.
We measured the root distribution of Hordeum vulgare (barley) and Beta vulgaris (sugar beet), grown in a glasshouse in cylindrical
containers with a volume of 1.3 L (length 26cm, inner diameter 8 cm). Measurements were done by non-destructive imaging of roots
using nuclear magnetic resonance imaging (MRI) as described extensively by Jahnke et al.(2009). This method is able to detect roots
with a diameter down to 300 mM, which implies that ne roots go unnoticed. Image segmentation of the root system was done by
thresholding the image and removing the stem and the sugar beet, with the RegionGrowingMacro module in MeVisLab (ver. 2.2.1;
MeVisLab, Bremen, Germany). After segmentation, the pixel values were integrated for the inner and outer regions of the soil core and
divided by the integral over the whole pot.
850 Functional Plant Biology H. Poorter et al.
... Some studies have been done on highbush blueberry production using containers, evaluating optimal pot size for greenhouse production [8,20] and estimating the impact on the substrate microclimatic conditions, metabolites content, and yield [8,21], while some others have focused on physical properties and substrate water distribution [22], as well as on analyzing the effects of rooting volume [23]. It is been suggested that reduced growth in smaller pots is caused mainly by a reduction in photosynthesis per unit leaf area, rather than by changes in leaf morphology or biomass allocation [23]. ...
... Some studies have been done on highbush blueberry production using containers, evaluating optimal pot size for greenhouse production [8,20] and estimating the impact on the substrate microclimatic conditions, metabolites content, and yield [8,21], while some others have focused on physical properties and substrate water distribution [22], as well as on analyzing the effects of rooting volume [23]. It is been suggested that reduced growth in smaller pots is caused mainly by a reduction in photosynthesis per unit leaf area, rather than by changes in leaf morphology or biomass allocation [23]. Photosynthesis of the blueberry cultivars used in this study grown in containers has not been studied before under Alabama conditions. ...
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Recently, there has been increased interest in container blueberry production as a viable alternative to open-field blueberry planting. Container production of blueberries offers numerous advantages, among these, a lack of limitation by suboptimal soil conditions in the open field and the ability to control substrate pH, drainage, and organic matter. The photosynthetic response for three container-grown Southern highbush blueberry (interspecific Vaccinium hybrids) cultivars including ‘Jewel’, ‘Meadowlark’, and ‘Victoria’ and a rabbiteye blueberry (Vaccinium virgatum) ‘Baldwin’, were measured during the spring and summer of 2022. It was hypothesized that the three cultivars evaluated would have different photosynthetic responses. The objective of this study was to determine the photosynthetic activity of different blueberry cultivars during the first year of crop establishment. A series of measurements were conducted every 2 h throughout the day and for different dates using a gas exchange data analyzer on newly matured fully expanded leaves located in the top middle section of the canopy for each cultivar. The response curves showed that net photosynthesis (A) became saturated at moderate light, with saturation occurring at a photosynthetic photon flux density (PPFD) of 1932 µmol m−2 s−1. At this point, the rate of CO2 assimilation was approximately 16.84 µmol CO2 m−2 s−1. No differences in (A) were found among cultivars. Overall, the attained values of photosynthesis provide a strong conceptual basis for understanding the cultivar variation response when grown in containers; therefore, the containerized system may serve as a production system for early fruiting blueberries in Alabama, USA.
... To determine the nutritional value of the plant, it is necessary to measure the amount of mineral elements. Measuring nutrients in plant tissue can be used as a tool to detect plant response to growth conditions as well as changes in growth and physiological indicators in plants 53 . Light causes changes in enzyme activities in the plant by influencing the production pathways of primary metabolites and thus affects the concentration of elements inside the plant 12 . ...
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The aim of this study was to investigate the impact of different cultivation systems (soil cultivation, hydroponic cultivation in greenhouse conditions, and hydroponic vertical cultivation in plant factory under different LED lights) and foliar spraying of nano calcium carbonate on pennyroyal plants. Nano calcium carbonate was applied to the plants at a 7-day interval, three times, one month after planting. Results showed that the greenhouse cultivation system with calcium carbonate foliar spraying produced the highest amount of shoot and root fresh mass in plants. Additionally, foliar spraying of calcium carbonate increased internode length and leaf area in various cultivation systems. Comparing the effects of different light spectrums revealed that red light increased internode length while decreasing leaf length, leaf area, and plant carotenoids. Blue light, on the other hand, increased the leaf area and root length of the plants. The hydroponic greenhouse cultivation system produced plants with the highest levels of chlorophyll, carotenoids, and phenolic compounds. White light-treated plants had less iron and calcium than those exposed to other light spectrums. In conclusion, pennyroyal plants grown in greenhouses or fields had better growth than those grown in plant factories under different light spectrums. Furthermore, the calcium foliar application improved the physiological and biochemical properties of the plants in all the studied systems.
... Applying results from pre-breeding to practical breeding depends on how well genotypes predict intended outcomes, like yield, under field conditions. Yet, there are significant differences between controlled and field conditions in the target environments (Poorter et al., 2012) for example regarding light intensities or room for root expansion. It is impossible to fully simulate outdoor environmental conditions in experimental setups due to their complex dynamics (Kumar et al., 2015). ...
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Mission-oriented governance of research focuses on inspirational, yet attainable goals and targets the sustainable development goals through innovation pathways. We disentangle its implications for plant breeding research and thus impacting the sustainability transformation of agricultural systems, as it requires improved crop varieties and management practices. Speedy success in plant breeding is vital to lower the use of chemical fertilizers and pesticides, increase crop resilience to climate stresses and reduce postharvest losses. A key question is how this success may come about? So far plant breeding research has ignored wider social systems feedbacks, but governance also failed to deliver a set of systemic breeding goals providing directionality and organization to research policy of the same. To address these challenges, we propose a heuristic illustrating the core elements needed for governing plant breeding research: Genetics, Environment, Management and Social system (GxExMxS) are the core elements for defining directions for future breeding. We illustrate this based on historic cases in context of current developments in plant phenotyping technologies and derive implications for governing research infrastructures and breeding programs. As part of mission-oriented governance we deem long-term investments into human resources and experimental set-ups for agricultural systems necessary to ensure a symbiotic relationship for private and public breeding actors and recommend fostering collaboration between social and natural sciences for working towards transdisciplinary collaboration.
... We had no measurements during this early period, but the model simulated this expected trend well and had the correct fRMF at 28 DAP. After 28 days, the measured fRMF was declining, as is generally true for older (Henn and Damschen 2021) and larger (Poorter et al. 2012) plants. The model predicted this decline quite well, especially when roots were classi ed based on diameter, as is done in the experimental data (Fig. 3E, blue line). ...
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· Background and Aims: Plants store carbohydrates for later use during, e.g., night, drought, and recovery after stress. Carbon allocation presents the plant with tradeoffs, notably between growth and storage. We asked how this tradeoff works for cassava (Manihot esculenta)pre- and post-storage root (SR) formation and if manipulation of the number of storage organs and leaf growth rate might increase yield. · Methods: We developed a functional-structural plant model, called MeOSR, to simulate carbon partitioning underlying cassava growth and SR formation in conjunction with the root system's three-dimensional (3D) architecture (RSA). We validated the model against experimental data and simulated phenotypes varying in the number of SR and leaf growth rate. · Results: The simulated 3D RSA and the root mass closely represented those of field-grown plants. The model simulated root growth and associated carbon allocation across development stages. Substantial accumulation of non-structural carbohydrates (NSC) preceded SR formation, suggesting sink-limited growth. SR mass and canopy photosynthesis might be increased by both increasing the number of SR and the leaf growth rate. · Conclusion: MeOSR offers a valuable tool for simulating plant growth, its associated carbon economy, and 3D RSA over time. In the first month, the specific root length increased due to root branching, but in the third month, it decreased due to secondary root growth. The accumulation of NSC might initiate SR development in cassava. Cassava growth is relatively slow during the first 3 months, and a faster crop establishment combined with a greater SR growth might increase yield.
... This usually involves shifted mean levels and much lower amplitudes and dynamics of the environmental variables than in the field, such as light intensities and air and soil temperatures due to technical limitations or cost-efficiency 11 . Furthermore, in most indoor systems, plant performance is limited by pot size constraints 13 . Thus, natural outdoor environments are more relevant for assessing the expression of performance traits in crops since phenotypes measured on plants cultivated in greenhouses or climate chambers often deviate greatly from those observed for plants exposed to natural conditions in open fields, even considering common agricultural management practices. ...
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In plant science, the suboptimal match of growing conditions hampers the transfer of knowledge from controlled environments in glasshouses or climate chambers to field environments. Here we present the PhenoSphere, a plant cultivation infrastructure designed to simulate field-like environments in a reproducible manner. To benchmark the PhenoSphere, the effects on plant growth of weather conditions of a single maize growing season and of an averaged season over three years are compared to those of a standard glasshouse and of four years of field trials. The single season simulation proves superior to the glasshouse and the averaged season in the PhenoSphere: The simulated weather regime of the single season triggers plant growth and development progression very similar to that observed in the field. Hence, the PhenoSphere enables detailed analyses of performance-related trait expression and causal biological mechanisms in plant populations exposed to weather conditions of current and anticipated future climate scenarios.
... Further investigations into nitrogen uptake rate, nitrogen use efficiency, total nitrogen content of plants, as well as the composition and abundance of AMF in the pots, are necessary. Furthermore, our previous studies (Liu et al., 2018; suggest that nitrogen availability rather than pot size likely constrained the plant growth, although we cannot rule out the possibility that some plants may have been root-bound, potentially influencing their responses to nitrogen fluctuations and AMF inoculation (Poorter et al., 2012). ...
Both enemies and mutualists play crucial roles in shaping plant invasion processes. Recent studies have suggested that resource fluctuations could indirectly promote plant invasion through higher trophic levels, such as enemies. However, the influence of mutualists like arbuscular mycorrhizal fungi (AMF) on plant invasion under nitrogen fluctuations remains untested. We conducted a pot mesocosm experiment using a three‐factorial experimental design to assess the individual and interactive effects of nitrogen availability, nitrogen fluctuation and AMF on invasive success of alien plants. We grew nine invasive alien species alongside five different native communities in pot mesocosms. These were then subjected to varied nitrogen availabilities (low vs. high), nitrogen fluctuations (constant vs. pulsed) and AMF presence or absence within a sterile substrate. We found that pulsed nitrogen supply increased the dominance of invasive alien species in low nitrogen availability, regardless of the presence or absence of AMF inoculation. However, in high nitrogen availability, pulsed nitrogen supply only enhanced this dominance in pots without AMF inoculation. This was tentatively evidenced by the three‐way interaction among nitrogen‐availability, nitrogen‐fluctuation and AMF‐inoculation treatments. Furthermore, the dominance promotion by nitrogen addition was greater than that by AMF inoculation. Synthesis and applications . Our findings present, for the first time, evidence that AMF may play a crucial role in mediating the promotion effects of nitrogen fluctuations on alien plant invasion. To better understand the invasion process of alien plants and evaluate their impact on native communities, future research should integrate abiotic and biotic drivers into a single framework. Furthermore, our findings underscore the importance of prioritizing habitats with higher nutrient availability and variability for protection against alien plant invasions.
... In future, some features of the phenotyping installation need to be taken into account when it is used to assess genotypic difference in responses to drought: due to the relatively large volume of the rhizo-pots compared to regular growth pots, soil moisture levels will decrease more slowly. This is closer to the natural drought scenarios in the field, where gradual changes in water availability occur rather than abrupt changes (Poorter et al., 2012). Soil drying in the rhizopots occurs due to the water consumption by the plants and the evaporation from the soil surface. ...
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In recent years, various automated methods for plant phenotyping addressing roots or shoots have been developed and corresponding platforms have been established to meet the diverse requirements of plant research and breeding. However, most platforms are only either able to phenotype shoots or roots of plants but not both simultaneously. This substantially limits the opportunities offered by a joint assessment of the growth and development dynamics of both organ systems, which are highly interdependent. In order to overcome these limitations, a root phenotyping installation was integrated into an existing automated non-invasive high-throughput shoot phenotyping platform. Thus, the amended platform is now capable of conducting high-throughput phenotyping at the whole-plant level, and it was used to assess the vegetative root and shoot growth dynamics of five maize inbred lines and four hybrids thereof, as well as the responses of five inbred lines to progressive drought stress. The results showed that hybrid vigour (heterosis) occurred simultaneously in roots and shoots and was detectable as early as 4 days after transplanting (4 DAT; i.e., 8 days after seed imbibition) for estimated plant height (EPH), total root length (TRL), and total root volume (TRV). On the other hand, growth dynamics responses to progressive drought were different in roots and shoots. While TRV was significantly reduced 10 days after the onset of the water deficit treatment, the estimated shoot biovolume was significantly reduced about 6 days later, and EPH showed a significant decrease even 2 days later (8 days later than TRV) compared with the control treatment. In contrast to TRV, TRL initially increased in the water deficit period and decreased much later (not earlier than 16 days after the start of the water deficit treatment) compared with the well-watered plants. This may indicate an initial response of the plants to water deficit by forming longer but thinner roots before growth was inhibited by the overall water deficit. The magnitude and the dynamics of the responses were genotype-dependent, as well as under the influence of the water consumption, which was related to plant size.
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Accurate estimates of current and future habitat suitability are needed for species that may require assistance in tracking a shifting climate. Standard species distribution models (SDMs) based on occurrence data are the most common approach for evaluating climatic suitability, but these may suffer from inaccuracies stemming from disequilibrium dynamics and/or an inability to identify suitable climate regions that have no analogues within the current range. An alternative approach is to test performance with experimental introductions, and model suitability from the empirical results. We used this method with the Haleakalā silversword (Argyroxiphium sandwicense subsp. macrocephalum), using a network of out-plant plots across the top of Haleakalā volcano, Hawaiʻi. Over a ~ 5-year period, survival varied strongly across this network and was effectively explained by a simple model including mean rainfall and air temperature. We then applied this model to estimate current climatic suitability for restoration or translocation activities, to define trends in suitability over the past three decades, and to project future suitability through 2051. This empirical approach indicated that much of the current range has low suitability for long-term successful restoration, but also identified areas of high climatic suitability in a region where plants do not currently occur. These patterns contrast strongly with projections obtained with a standard SDM, which predicted continued suitability throughout the current range. Under continued climatic shifts, these results caution against the common SDM presumption of equilibrium between species’ distributions and their environment, even for long-established native species.
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Microplastics can affect their surroundings physically and chemically, resulting in diverse effects on plant-soil systems. Similar to other substances (e.g. nutrients and water), microplastics in the environment occur in patches. Such heterogeneous distributions could affect plant responses to plastic pollution. Yet, this has remained untested. We conducted a multispecies experiment including 29 herbaceous plant species and three different microplastic treatments (a control without microplastics, a homogeneous and a heterogeneous microplastic distribution). Based on biomass and root-morphological traits, we assessed how different plastic distributions affect the performance and root-foraging behavior of plants, and whether stronger root foraging is beneficial when microplastics are distributed patchily. Next to general effects on plant productivity and root morphology, we found very strong evidence for root-foraging responses to patchy plastic distributions, with a clear preference for plastic-free patches, resulting in 25% longer roots and 20% more root biomass in the plastic-free patches. Interestingly, however, these foraging responses were correlated with a reduced plant performance, indicating that the benefits of plastic avoidance did not compensate for the associated investments. Our results provide new insights in plant-microplastic interactions and suggest that plants might not just be passively affected by but could also actively respond to environmental plastic pollution.
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Background Suboptimal nitrogen availability is a primary constraint for crop production in low-input agroecosystems, while nitrogen fertilization is a primary contributor to the energy, economic, and environmental costs of crop production in high-input agroecosystems. In this article we consider avenues to develop crops with improved nitrogen capture and reduced requirement for nitrogen fertilizer. Scope Intraspecific variation for an array of root phenotypes has been associated with improved nitrogen capture in cereal crops, including architectural phenotypes that colocalize root foraging with nitrogen availability in the soil; anatomical phenotypes that reduce the metabolic costs of soil exploration, improve penetration of hard soil, and exploit the rhizosphere; subcellular phenotypes that reduce the nitrogen requirement of plant tissue; molecular phenotypes exhibiting optimized nitrate uptake kinetics; and rhizosphere phenotypes that optimize associations with the rhizosphere microbiome. For each of these topics we provide examples of root phenotypes which merit attention as potential selection targets for crop improvement. Several cross-cutting issues are addressed including the importance of soil hydrology and impedance, phenotypic plasticity, integrated phenotypes, in silico modeling, and breeding strategies using high throughput phenotyping for co-optimization of multiple phenes. Conclusions Substantial phenotypic variation exists in crop germplasm for an array of root phenotypes that improve nitrogen capture. Although this topic merits greater research attention than it currently receives, we have adequate understanding and tools to develop crops with improved nitrogen capture. Root phenotypes are underutilized yet attractive breeding targets for the development of the nitrogen efficient crops urgently needed in global agriculture.
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Growth of Magnolia grandiflora Hort. `St. Mary' (southern magnolia) trees in containers spaced 120 cm on center was studied for 2 years. During the 1st year, trees were grown in container volumes of 10, 27, or 57 liter. At the start of the second growing season, trees were transplanted according to six container shifting treatments [10-liter containers (LC) both years, 10 to 27LC, 10 to 57LC, 27LC both years, 27 to 57LC, or 57LC both years]. The mean maximum temperature at the center location was 4.8 and 6.3C lower for the 57LC than for the 27 and 10LC, respectively. Height and caliper, measured at the end of 2 years, were” greatest for magnolias grown continuously in 27 or 57LC. Caliper was greater for trees shifted from 10LC to the larger containers compared with trees grown in 10LC both years. Trees grown in 10LC both years tended to have fewer roots growing in tbe outer 4 cm of the growing medium at the eastern, southern, and western exposures. During June and August of the 2nd year, high air and growth medium temperatures may have been limiting factors to carbon assimilation. Maintenance of adequate carbon assimilation fluxes and tree growth, when container walls are exposed to solar radiation, may require increasing the container volume. This procedure may be more important when daily maximum air temperatures are lower during late spring or early fall than in midsummer, because low solar angles insolate part of the container surface.
Seedling pecan top growth was greater in 38 1(#10) containers and shallow 19 1(#5) containers when compared to deeper 19 1 (#5) containers and 11 1 (#3) containers; however, all trees were large enough to bud by July of the first growing season. Root pruning at transplanting did not affect top growth, but increased root branching and total root growth. Increased rates of a complete fertilizer increased root growth, but did not affect top growth.
Leyland cypress (X Cupressocyparis leylandii ‘Haggerston Grey’) cuttings were grown in 3.8 1 (#1), 7.6 1 (#2) and 11.4 1 (#3) containers in media combinations of pine bark, hardwood bark and coarse sand. Plants were fertilized with weekly applications of 200, 400, 600 or 800 ppm N. A growth index, percent foliar nitrogen, pH and soluble salts were detennined after 1 growing season and top dry weight was measured after 2 growing seasons. The 4 pine bark: 1 sand (v/v) and combination medium had lower pH values than the 4 hardwood bark: 1 sand medium. Percent foliar nitrogen ranged from 0.9% N to 1.34% N. Incremental N application did not increase N accumulation in leaf tissue. Increasing container size or N application did not increase the growth index through the fIrst growIng season. By the end of the second growing season plants in 11.4 1 containers were larger than those in 3.8 1 containers.
Transplants for both vegetable and floral crops are produced in a number of various sized containers or cells. Varying container size alters the rooting volume of the plants, which can greatly affect plant growth. Container size is important to transplant producers as they seek to optimize production space. Transplant consumers are interested in container size as it relates to optimum post-transplant performance. The following is a comprehensive review of literature on container size, root restriction, and plant growth, along with suggestions for future research and concern.
Many experiments are conducted in greenhouses or growth chambers in which plants are grown in pots. Considerable research has shown that pots can have a limiting effect on overall plant growth. This research was undertaken to examine the effects of pot size specifically on transpiration response of maize (Zea mays L.) and soybean (Glycine max L.) plants undergoing water-deficit stress. Maize and soybean experiments were conducted similarly, but as separate experiments. Maize plants were grown in 2.3, 4.1, 9.1, and 16.2 I pots sealed to prevent water loss except by transpiration. For each pot size, plants were divided into two watering regimes, a well-watered control and a water-deficit regime. Water deficits were imposed by simply not rewatering the pots. Soybean was examined in a similar manner, but only the three larger pot sizes were used in the experiment. For both maize and soybean, and in both watering regimes, there was a significant reduction of shoot dry weight and total transpiration with decreasing pot size. However, there were no significant differences among pot sizes in the fraction of transpirable soil water (FTSW) point at which transpiration began to decline (FTSW≃0.31 for maize and ≃0.35 for soybean) or in the overall relationship of transpiration rate to soil water content in response to water deficits. These results indicated that, regardless of pot size or plant size, the overriding factor determining transpirational response to drought stress was soil water content.
The wax-apple [Syzygium samarangense (BI.) Merr. and Perry] is a vigorous tropical fruit tree species that has five to six growth flushes per year. One-year-old, root-bearing wax-apple trees were grown in different-sized containers filled with potting mixture to test if container volume restricts shoot and/or root growth and thereby lends itself to forcing culture. The trunk cross-sectional area (TCSA) at 15 cm above the soil was measured to assess vegetative growth. After 6 months, the TCSA had increased quadratically with container volume. At the end of the first and second year, leaf count, leaf area, leaf dry mass, stem dry mass, shoot dry mass, and root dry mass were positively correlated with container volume. However, the shoot: root ratios remained fairly constant among treatments during the experimental period. Thus, root restriction is an effective means of reducing shoot and root growth of the wax-apple.
Publisher Summary This chapter explains the growth and behavior pattern of roots and shoots And discusses the generation of hormone like messages due to environmental changes in detail. Hormonal messages are recycled between roots and shoots. Roots import hormones and shoots acts as active hormone sinks. Various techniques to measure hormone levels and activities are described, the primary being the samples taken from the sap. Hormone traffic between roots and shoots forms the basis for the morphological and functional control that roots can exert on shoots. Regulation of root: shoot ratio by roots with mineral supply is explained in detail. Hormone like action on roots is responsible for regulation of protein levels in leaves and flowering in plants. The influence of hormone like messages by roots for development of shoots is presented. The influence of plant hormones cytokinins, gibberellins, abscisic acid, auxin and ethylene are explained. Cytokinins are main source of hormones in plants. They are carried by transpiration from the root tips present on a root system to recipient shoot tissues. Gibberellins are important in delaying leaf senescence. The output of gibberellins under stress is discussed. Abscisic acid is an important regulator of stomatal closure. Ethylene–mediated responses in the shoot are explained.
The two main proximate causes of intratree variation in individual seed mass of lodgepole pine [Pinus contorta var. latifolia (Dougl. ex Loudon)] were space constraints within cones and between-branch position effects. Cone scales get smaller near the distal tip of the cone. Seed predation has selected for the production of the fertile zone in the distal tip of the cone where cone scales are smaller. Thus, seed and wing size were constrained by cone scale size, and decreased from the base to the tip of the cone. Heavier seeds had larger seed wings, but wing loading (fruit mass/wing area) was independent of seed mass. Although it is possible that seed mass variation might be selected because it increases dispersal effectiveness, several factors - the consistent within-cone positional variation in seed mass, wing area, and wing loading; the dependence of seed mass, wing area, and wing loading on cone size; and the positive relationship between seed mass and wing area - suggest that variation is caused by constraints by scale size.