Scientia Horticulturae 105 (2005) 269–282
www.elsevier.com/locate/scihorti
Nutrient solution effects on the development
and yield of Anthurium andreanum Lind.
in tropical soilless conditions
L. Dufour a,1,*, V. Guérin b
a
Unité de Recherche AgroPédoclimatologique de la zone Caraı̈be, INRA Antilles-Guyane,
Domaine Duclos, 97170 Petit-Bourg, France
b
UMR A_462 SAGAH (INRA/INH/Univ. d’Angers), B.P. 60057,
49071 Beaucouzé Cedex, France
Received 12 November 2002; received in revised form 21 January 2005; accepted 21 January 2005
Abstract
The fertilization of anthurium grown in soilless culture in tropical countries is often empirically
based. The methods used generally lead the grower to overestimate plant needs and to apply
excessive quantities of nutrients. Mineral elements, and thus money, are wasted and there is a risk
of pollution of groundwater and watercourses. In order to improve our knowledge of plant
requirements, we measured, over 2 1/2 years, the growth and yield of anthurium plants receiving
nutrient solutions with different total nitrogen, potassium and calcium concentrations and different
NH4+/NO3 ratios. Mineral analyses of plant parts, of nutrient, leachate and substrate solutions
and of the solid substrate were carried out throughout plant development. Plants receiving
4.5 mmol N/l and 1.6 mmol K/l in the nutrient solution had significantly slower growth and
lower yield compared to those receiving 8.9 mmol N/l and 3.2 mmol K/l. For these latter N and K
concentrations, a N–NH4+/N–NO3 ratio of 0.37 and a calcium concentration of 1.15 mmol/l gave
better plant growth, development and yield than a ratio of 0.24 and a calcium concentration of
2.25 mmol/l. Applying the nutrient solution containing 8.9 mmol N/l and 3.2 mmol K/l with a
N–NH4+/N–NO3 ratio of 0.37 resulted in a shorter vegetative period and more and larger flower
production. The calculated mineral balances of the crop showed that more than 60% of the
supplied nutrients were lost in the leachate. Suitable nutrient solutions are proposed in order to
* Corresponding author. Tel.: +33 4 67 61 75 71; fax: +33 4 67 61 55 12.
E-mail address: [email protected] (L. Dufour).
1
Permanent address: UMR-System (INRA/CIRAD/ENSAM), CIRAD, Avenue Agropolis, TA 40/01, Bât. 1,
34398 Montpellier Cedex 5, France.
0304-4238/$ – see front matter # 2005 Elsevier B.V. All rights reserved.
doi:10.1016/j.scienta.2005.01.022
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L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
match plant absorption at different crop growth stages. The volume of nutrient solution supplied
can be reduced to limit the amount of leachate, but as water demand is high, there must be at least
30% of leaching to avoid salt accumulation in the substrate. Adjusting the nutrient solution volume
and composition to match plant requirements is the first step for flower yield improvement,
fertilizer efficiency and reduction of pollution.
# 2005 Elsevier B.V. All rights reserved.
Keywords: Anthurium andreanum Lind.; Mineral requirements; Mineral analyses; Plant growth; Cut flower
production; Soilless cultivation
1. Introduction
Soilless cultivation of anthurium was developed in the French West Indies in the 1980s
because of the unintentional introduction of two bacterial diseases and the market demand
for better flower quality. This cultivation technique is frequently achieved using an
automatic fertigation system. The volume of substrate available to roots is restricted and
the material used generally has a low cation exchange capacity. Plant nutritional status
affects yield, quality, and susceptibility to bacterial (Pérez et al., 1992) and fungal diseases
(Sakaı̈, 1990). Hence it is essential to have a good knowledge of the plant’s mineral
requirements in order to ensure a good yield and to avoid nutrient wastage. The matching of
supply and demand may decrease production costs and reduce the risk of water pollution.
On the other hand, it is necessary to limit mineral imbalance in the medium by assuring a
minimal leaching of 30% (Coı̈c and Lesaint, 1983; Roeber, 1999) by supplying excess
nutrient solution.
We are interested in knowing the influence of different fertilization patterns on
anthurium growth and development. Experiments on this species (Boertje, 1978; Dufour
and Clairon, 1997; Sonneveld and Voogt, 1993) showed that its nitrogen supply must not
exceed 6 g of nitrogen per plant per year, otherwise flower quality declines. These authors
also emphasized the importance of potassium supply on flower production and flower
quality. Higaki et al. (1992) showed that lack of K has a big influence on flower stem length.
However, nutrient uptake can be affected by factors such as environmental conditions,
irrigation, type of fertilizer used and methods of application. Therefore, fertilization studies
must be supplemented by (i) following the mineral elements’ availability in relation to
changes in the ionic composition of the substrate, which can be determined using induced
percolate analysis (Lemaire et al., 1995; Marfá et al., 2002); (ii) assessing plant nutrient
absorption by leaf mineral analysis (Boertje, 1978; Higaki et al., 1980, 1992; Mills and
Scoggins, 1998).
These two parameters were measured in relation to nutrient supply. The results of plant
and solution analysis allowed us to establish a crop nutrient balance in order to have a better
knowledge of culture parameters and to estimate the fertilizer supply efficiency.
In this study, we observed the growth and development of anthurium plants with three
different fertilization formulations. We chose to study the variation in calcium
concentration because of the influence of this nutrient on flower quality shown by
Higaki et al. (1980), and of nitrogen because of the pollution caused by nitrate leaching.
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271
Finally, the estimation of plant nutrient uptake allowed us to propose a fertilization pattern
for each developmental phase.
2. Materials and methods
2.1. Plant materials and cultivation conditions
The experiment took place at the Duclos experimental station of INRA Antilles-Guyane
(16870 N, 618240 W, 110 m above MSL). The environment is characterized by a warm humid
tropical climate with an annual average temperature of 24.6 8C, annual rainfall of 2800 mm
and PAR (photosynthetically active radiation) values of 27 to 41 mol/m2/day.
Tissue-cultured plantlets of Anthurium andreanum ‘Cancan’ (Anthura B.V. variety)
were planted in 2.5 l pots filled with a substrate consisting of 5–15 mm pouzzolane (local
volcanic gravel) and composted and disinfected wood chips (2:1, v/v). At planting, the
plantlets measured 8–10 cm from the base to the tip of the largest leaf. One year after
planting, the plants were repotted into 5 l pots, with the same substrate. The planting
density was around 5/m2 of greenhouse. The pots were grown in a shadehouse whose
cladding materials excluded 87% of the PAR.
Three nutrient solutions were supplied automatically by trickle irrigation. Each plant
received approximately 130 ml of nutrient solution per day for 1 1/2 years and 200 ml,
thereafter. Four replications of 64 plants each per treatment were sited in the shadehouse in
order to allow for spatial variation (e.g. of microclimate). Treatments and replications were
laid out in an experimental design of four beds according to a Youden scheme. The
composition of the nutrient solutions is given in Table 1. The solution called N:K:Ca
(1:1:1) was the one that gave the best yields in Dufour and Clairon’s (1997) experiments:
we regarded it as the control treatment. In the N:K:Ca (1:1:0.5) solution, we wanted to test
if a 50% decrease in calcium supply has a negative effect on flower quality, as stated by
Higaki et al. (1980). The decrease in calcium cations in this solution was partly balanced by
the increase in ammonium cations. Lastly, in order to reduce nitrate leaching, we decreased
the nitrogen concentration by 50% in the N:K:Ca (0.5:0.5:0.5) solution, keeping the same
weight ratios between N, K and Ca (1:1:0.7). The three solutions had the same phosphorus,
magnesium, iron and micronutrient concentrations.
The fertilizers used to make these solutions were monoammonium phosphate,
magnesium sulfate, nitric acid, calcium nitrate (this fertilizer contains 1.4 g of ammonium
for 100 g) and potassium nitrate; calcium chloride for the N:K:Ca (1:1:1) and N:K:Ca
Table 1
Composition of the nutrient solutions (mmol/l)
Treatment N:K:Ca
N–NO3
N–NH4
Total N
H2PO4
K
Ca
Mg
1:1:1
1:1:0.5
0.5:0.5:0.5
7.2
6.5
2.9
1.8
2.4
1.5
8.9
8.9
4.5
1.4
1.4
1.4
3.2
3.2
1.6
2.3
1.2
1.2
1.6
1.6
1.6
The three solutions had the same Fe concentration: 1.3 mmol/l, and micronutrients content (in mmol/l): ZnSO4:
5.6; MnSO4: 10.7; CuSO4: 1.5; H3BO3: 18.8; NH4Mo: 3.4.
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(0.5:0.5:0.5) solutions; ammonium nitrate for the N:K:Ca (1:1:0.5) solution and potassium
chloride for the N:K:Ca (0.5:0.5:0.5) solution.
2.2. Plant growth measurements
The 10 plants in the middle of each replication were measured. In order to quantify their
growth, the youngest leaf and the growing flower (when present) of these plants were
measured once a week: petiole and blade length, blade width, peduncle, spathe and spadix
length and spathe width. The leaf area during the reproductive phase was estimated from
the equation (Dufour and Guérin, 2003):
Leaf area ¼ 0:92 blade length blade width
On a single day, we cut and counted the mature flowers in the whole shadehouse, to
calculate the yield for each treatment.
2.3. Mineral analysis
2.3.1. Plant analysis
Two plants per replication were removed at five growth stages: during vegetative growth
(5 months after planting), at first flower emergence (7 months after planting), at first flower
harvest (9 months after planting), at large flower appearance (11 months after planting) and
during full production (27 months after planting, end of the experiment). The leaves and
flowers of these plants were measured, the dry weights of their different parts (roots, mature
leaf blades, young leaf blades, mature leaf petioles, young leaf petioles, peduncle, spathe,
spadix and stem) were recorded. These parts were ground and their mineral composition
(N, P, K, Ca and Mg) was analyzed after mineralisation following the technique described
by Novosamsky et al. (1983).
Plant nutrient uptake was calculated from the results of plant analyses, adding an
assessment of the losses due to the leaf pruning and flower harvest.
2.3.2. Solution analysis
Six times during the experiment, i.e. during the vegetative phase (5 months after
planting); when the first flower developed (6 months after planting); at the beginning of
flower production (8, 10 and 12 months after planting) and during full production (17
months after planting), nutrient solution, leachate and substrate solution were collected and
analyzed. Percolates were collected from the bottom of the containers after supplying
about 150 ml of nutrient solution below the dripper. As shown by Lemaire et al. (1995), the
percolate composition gives a good assessment of the substrate solution composition
(substrate at saturation).
Electrical conductivity (EC), pH, NO3–N, NH4+–N, P, K, Ca, and Mg concentrations
of all the solutions were determined.
Supply and leaching losses were the total amounts of mineral elements applied to the
plant and drained from the pot, respectively. They were calculated from the nutrient and
leachate solution compositions multiplied by their volumes. Substrate solution
concentration was estimated by variation in at the two considered dates.
L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
273
2.3.3. Substrate analysis
In order to check the nutrient balances, we took substrate samples for analysis three
times during the experiment: at the end of the vegetative phase, at the beginning of the
reproductive phase (just before repotting) and at the end of the experiment, with mature
plants. The analyses were performed as described by Marfá et al. (2002).
3. Results
3.1. Influence of fertilization on plant development and yield
Anthurium plantlet growth begins with a vegetative period during which no flowers are
produced (Dufour and Guérin, 2003). The duration of this period was strongly influenced
by fertilization (Fig. 1); analysis of variance shows a significant difference between the
treatments. Plants of the N:K:Ca (1:1:0.5) treatment produced their first flower
approximately 7 months after planting, more than half a month before plants of the
N:K:Ca (1:1:1) treatment and more than a month before those of the N:K:Ca (0.5:0.5:0.5)
treatment. The plants that received the solution with the lowest total N concentration
(4.5 mmol/l) had a longer vegetative phase than the others. The N:K:Ca (1:1:0.5) solution
contained half the calcium of N:K:Ca (1:1:1), but it also had a higher ammonium fraction
for the same total amount of N (Table 1).
The number of harvested flowers (Fig. 1), the leaf size (Fig. 2a) and the quality of
harvested flowers (Fig. 2b) were also influenced by the nutrient solution. The plants
supplied with the N:K:Ca (1:1:0.5) solution produced significantly more and larger flowers
and had larger leaves than the others. The plants in this treatment produced 6.4 0.4
(standard deviation) flowers/plant during the second year of production. In the N:K:Ca
(1:1:1) and N:K:Ca (0.5:0.5:0.5) treatments, the plants produced 5.4 0.1 and 5.5 0.3
Fig. 1. Length of vegetative period (cross-hatched columns), as the number of days between planting and
appearance of the first flower bud; means for the 40 observed plants supplied with the same nutrient solution.
Number of flowers harvested (gray columns) per plant during the experiment; means of all the plants supplied with
the same nutrient solution. Vertical bars indicate Student’s t-test intervals at the 5% probability level.
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Fig. 2. Leaf area (a) and flower size (b) development during the reproductive period; means of the 40 measured
plants per treatment (squares for the N:Ca:K (1:1:1) treatment, triangles for the N:Ca:K (1:1:0.5) treatment, circles
for the N:Ca:K (0.5:0.5:0.5) treatment). Vertical bars represent Student’s t-test intervals at the 5% probability
level.
flowers/plant, respectively, during the same period. The apparent phyllochron is the time
between the emergence of two leaves. Since a flower is produced for each leaf, the apparent
phyllochron gives a good representation of the flower growth rate. The plants of the N:K:Ca
(1:1:0.5) treatment had a shorter apparent phyllochron than the others (Fig. 3). The
differences from the N:K:Ca (0.5:0.5:0.5) treatment are significant except for leaves 8
and 9. The only significant differences from the N:K:Ca (1:1:1) treatment are for leaves
11 and 12.
3.2. Mineral analyses
The mineral element contents (mg/plant) of the entire plant in each treatment during the
whole experiment are given in Table 2. For the last two samplings, these data take into
account the plant parts removed by pruning and harvest. Significant differences were
observed for N:K:Ca (0.5:0.5:0.5) plant N content from the beginning of flower emergence
to the end of the experiment. These plants contained less N than all of the others. Twenty-
L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
275
Fig. 3. Apparent phyllochron during the reproductive phase. Vertical bars represent Student’s t-test intervals at the
5% probability level.
seven months after planting, the N:K:Ca (1:1:0.5) plants had absorbed significantly greater
amounts of all nutrients than the others. This was true even for P and Mg, whose solution
concentrations were the same as in the other treatments, and Ca, whose concentration was
lower in the N:K:Ca (1:1:0.5) than in the N:K:Ca (1:1:1) solution. These differences are
Table 2
Mineral uptake by the plants (mg/plant)
Montha
Solution N:K:Ca
5
1:1:1
1:1:0.5
0.5:0.5:0.5
46 a
53 a
37 a
10 a
11 a
10 a
88 a
87 a
77 a
48 a
42 a
40 a
27 a
25 a
24 a
7
1:1:1
1:1:0.5
0.5:0.5:0.5
194 b
331 a
121 b
37 b
69 a
37 b
277 b
481 a
221 b
210 b
283 a
176 b
95 b
154 a
99 b
N
P
K
Ca
Mg
First flowers harvest
9
1:1:1
1:1:0.5
0.5:0.5:0.5
249 b
377 a
109 c
53 b
83 a
36 b
347 b
549 a
194 c
305 a
360 a
174 b
128 b
198 a
95 b
11
1:1:1
1:1:0.5
0.5:0.5:0.5
502 a
542 a
198 b
112 a
103 a,b
79 b
683 a
758 a
333 b
623 a
495 b
346 c
261 a
256 a
214 a
27
1:1:1
1:1:0.5
0.5:0.5:0.5
1134 b
2368 a
660 c
1376 b
3589 a
788 c
1598 b
2093 a
1110 c
234 b
335 a
243 b
688 b
875 a
756 a,b
Mean of the eight analyzed plants per treatment—data were analyzed by analysis of variance, using Newman and
Keuls test at P = 0.05 for separation of means (inside a box). In the same box, data followed by the same letter are
not significantly different.
a
Month: number of months from planting to sampling.
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L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
Table 3
Mineral content of Anthurium andreanum leaves (g/100 g of dry matter)
Author
Leaves
N
P
K
Ca
Mg
Mills and Scoggins (1998)
N:K:Ca (1:1:1) treatment
N:K:Ca (1:1:0.5) treatment
N:K:Ca (0.5:0.5:0.5) treatment
Young
Young
Young
Young
2.6
1.8 b
2.8 a
1.8 b
0.4
0.3 a
0.3 a
0.4 a
3.0
2.4 b
3.1 a
2.2 b
1.2
1.8 a
1.6 a
1.6 a
0.6
0.8 b
0.5 c
1.2 a
Boertje (1978)
Higaki et al. (1992)
Higaki et al. (1980)
Mills and Scoggins (1998)
N:K:Ca (1:1:1) treatment
N:K:Ca (1:1:0.5) treatment
N:K:Ca (0.5:0.5:0.5) treatment
Last mature
Last mature
Mature
Mature
Mature
Mature
Mature
2.0
1.9
2.1
1.7
1.5 b
1.9 a
1.4 b
0.4
0.2
0.2
0.2
0.2 a
0.2 a
0.3 b
2.9
2.1
2.0
1.7
1.4 b
2.5 a
1.0 c
0.4
1.1
0.8
1.0
2.1 a
1.6 b
2.0 a
1.3
0.3
0.3
0.6
0.8 b
0.6 c
1.2 a
Several authors’ results compared with our results (means of the blade composition for the 24 plants sampled at the
end of the experiment). Comparisons between treatments were performed by analysis of variance, using Newman
and Keuls test at P = 0.05. In the same box, data followed by the same letter are not significantly different.
partly due to the higher dry weight of the N:K:Ca (1:1:0.5) plants (54 14.2 g versus
39.2 10.1 g and 31.0 10.1 g for N:K:Ca (1:1:1) and N:K:Ca (0.5:0.5:0.5) plants,
respectively). For every sampling date and almost every fertilization treatment (except
N:K:Ca (0.5:0.5:0.5) for the two last dates), potassium was the mineral element present in
the largest amount.
In Table 3, we compare the mineral composition of the leaf blades of plants 27 months
after planting with other authors’ results. For the N:K:Ca (1:1:0.5) solution, our N, P and K
concentrations were similar to those of other authors; the Mg concentration was lower and
Ca concentration much higher (by almost 1%) in mature blades. For the two other
solutions, N and K concentrations were lower and Ca was higher in our plants than in the
other authors’. Mg concentration was significantly lower in N:K:Ca (1:1:0.5) plant leaves,
while these plants’ uptake was higher (Table 2). The biomass of the N:K:Ca (1:1:0.5) plants
being higher than the others’, the Mg was more diluted in these plants. The same
observation can be made for Ca in mature leaves.
Table 4 shows nutrient use efficiency and leaching losses during the two developmental
phases of the plant. Potassium uptake by the plant was higher than that of the other
elements for all fertilization treatments and both growth stages. Nutrient use efficiency was
highest for the N:K:Ca (1:1:0.5) treatment during the vegetative phase—around 50%,
except for P, for which it was 17%. This may have been due to the faster growth of the
N:K:Ca (1:1:0.5) plants during this phase, which would have increased their nutrient
uptake. The plants of the N:K:Ca (1:1:1) treatment during the reproductive phase had
higher nutrient use efficiency (almost 45%, or 20% for P) than the others. The increase of
dry matter was a little faster for the N:K:Ca (1:1:1) plants than for the N:K:Ca (1:1:0.5)
ones during the reproductive period, which would explain their faster nutrient uptake.
The amount of nutrient elements lost by leaching in the N:K:Ca (1:1:0.5) treatment
varied between 82% (Ca in the vegetative phase) and 50% (P in the reproductive phase) of
the supply, and the leachate volume was on average 77% of the applied volume (data
not shown). This is much higher than the 30% recommended by Coı̈c and Lesaint (1983).
Element
Solution N:K:Ca
Vegetative phase
2
Supply (g/m )
Reproductive phase
Plant uptake
2
Leaching losses
2
(g/m )
(%)
(g/m )
(%)
Supply (g/m2)
Plant uptake
Leaching losses
(g/m2)
(%)
(g/m2)
(%)
N
1:1:1
1:1:0.5
0.5:0.5:0.5
3.35
3.12
1.98
0.63 b
1.25 a
0.35 b
18.6 y,z
41.0 x
19.3 x,y,z
1.78 a
1.96 a
0.73 b
53.5 x,y
63.4 x
37.4 y,z
5.84
6.18
4.07
2.04 l
1.40 l
0.51 m
35.3 x
23.1 x,y
12.3 x,y
4.05 l
3.74 l
1.08 m
69.5 x
61.7 x,y
26.6 z
P
1:1:1
1:1:0.5
0.5:0.5:0.5
1.10
1.46
1.36
0.10 b
0.24 a
0.12 a,b
9.4 y
17.1 x,y
8.5 y
0.67 b
0.86 a
0.83 a,b
60.9 y
58.9 x,y
60.2 x,y
2.55
2.84
2.83
0.50 m
0.23 l
0.28 lm
19.4 x
8.1 y
10.8 x,y
1.87 l
1.39 l
1.85 l
72.8 x
50.0 x,y
65.3 x,y
K
1:1:1
1:1:0.5
0.5:0.5:0.5
5.78
3.78
2.26
0.78 b
1.70 a
0.61 b
13.4 z
46.5 y
27.9 y,z
2.11 a,b
2.42 a
1.37 b
37.8 y
65.1 x
64.0 x,y
6.01
6.93
4.63
2.69 l
1.83 l,m
0.90 m
45.3 y
26.4 z
19.4 z
4.10 l
3.75 l,m
1.63 m
69.7 x
54.1 x,y
35.2 y
Ca
1:1:1
1:1:0.5
0.5:0.5:0.5
3.60
1.93
1.65
0.63 b
1.01 a
0.57 b
18.2 z
53.7 y
42.0 z,y
2.44 a
1.55 b
1.10 b
68.1 x
82.0 x
82.8 x
6.24
3.57
3.22
2.74 l
1.41 m
1.13 m
45.3 y
40.8 y
36.5 y,z
5.07 l
2.54 m
1.76 m
83.7 x
73.5 x
54.8 x
Planting density was 5.14 plants/m2. Means were compared at P = 0.05. Significant difference between solutions for one element in plant uptake or leaching losses is
indicated by different letters (a and b for vegetative; m and l for reproductive phase). Significant differences between the three solutions over both phases, for one element in
plant uptake % and leaching losses % are shown by letters x, y and z.
L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
Table 4
Nutrient supply, plant uptake and leaching losses during the first year of the experiment: in the vegetative phase (from 5 to 8 months after planting) and in the reproductive
phase (from 8 to 13 months after planting)
277
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L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
The nutrient supply always greatly exceeded plant absorption, even for the N:K:Ca
(0.5:0.5:0.5) treatment.
The variation of substrate solution nutrient content was very little (data not shown),
compared with uptake or leaching, except for nitrogen during the reproductive phase of
N:K:Ca (0.5:0.5:0.5) plants. In this case, the substrate solution nitrogen content decreased
by 0.98 g/m2 while uptake was 0.51 g N/m2. There was a high percentage of unaccountedfor N in this treatment. This N could have been lost due to NH3 volatilization or
denitrification losses from the substrate (Cabrera, 2003).
Substrate analysis (data not shown) revealed large variation between pots, especially for
water-soluble K content. However, we observed an increase in the phosphorus
concentration: the amount of water soluble P increased steadily from 1 mg/l of substrate
to 4 mg/l during the course of the experiment. This can be partly explained by the excessive
P supply in all the treatments: the sum of plant uptake and leaching losses was always less
than the supply. On the other hand, there was a substantial decrease in ammonium acetateextracted Ca during the growing period (around 1000 mg/l of new substrate versus 450 mg/
l of substrate at the end of the experiment for the N:K:Ca (1:1:0.5) treatment). As the sum
of plant Ca uptake and leaching losses was frequently higher than supply, it is possible that
the substrate gave up some of its Ca.
4. Discussion
4.1. Effects of total nitrogen and potassium supply on plant growth
Total nitrogen and potassium supply have an influence on the length of the juvenile
phase: with a lower level of supply, the plant produces its first flower later. N and K uptake
of the plants is always very much lower than supply, largely because of the small amount of
dry matter produced. All the leaves are smaller for the solution in which N and K are less
concentrated, yet they need more time to develop. Growth of leaves and flowers is slower.
When a plant with a poor nitrogen and potassium supply is in the full production phase, its
mature leaf blades contain 1.4% nitrogen and 1.0% potassium, which is less than for other
plants and the values found by Higaki et al. (1980) and Mills and Scoggins (1998). The
assimilate source, represented by the mature leaf blades, is consequently smaller (smaller
leaves with lower N and K concentrations). That may be why the sixth to ninth flowers are
smaller for the N:K:Ca (0.5:0.5:0.5) treatment. Before it has a positive net photosynthesis
rate, the young leaf is a metabolic sink which competes with the developing flower (Daı̈ and
Paull, 1990). The other flowers are not significantly smaller even when the leaves are, but
the plant produces fewer flowers when nitrogen is restricted.
4.2. Effects of N–NH4+/N–NO3 ratio and of calcium concentration
In our conditions, the N–NH4+/N–NO3 ratio also seems to affect plant growth and
development: when it increases from 0.24 to 0.37, while calcium concentration decreases
from 2.3 to 1.2 mmol/l, the juvenile phase is shortened and the leaves are larger.
The peduncle is longer for flowers 5–9, but the spathe size is not significantly different.
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279
The plant also produces more flowers when the ratio is higher. Claussen and Lenz (1999)
showed that the effects of nitrate and ammonium on growth reflect different plant species’
adaptability to soil conditions, particularly acidity. On the other hand, for a species quite
closely related to anthurium, Maranta leuconeura, N forms ranging from 5 to 75% of
ammonium do not influence plant growth (Strojny, 1999). van Herk et al. (1998) located the
optimum soil pH for Anthurium andreanum Lind. at around 5.7, and our substrate varied
from pH 8 at planting to 5.2 at the end of the experiment. This means that anthurium can
tolerate different pH conditions and its nitrogen assimilation is probably enhanced by the
presence of the two N forms.
4.3. Plant mineral composition
Considering the total quantity of mineral elements in the adult plants, the plants that
received the N:K:Ca (0.5:0.5:0.5) solution have approximately a third of the nitrogen
(350 mg of N/adult plant) found in the plants grown in the N:K:Ca (1:1:0.5) solution
(1000 mg of N/adult plant). Hence, it is likely that the N:K:Ca (0.5:0.5:0.5) plants’ reserves
are much lower and the life span of the crop should be lower. K and Ca contents were also
lower. The Ca assimilation seems to be enhanced when the growth is better; the total
amount of Ca in plants in the N:K:Ca (1:1:0.5) treatment being higher than for N:K:Ca
(1:1:1). But, as the dry weight of N:K:Ca (1:1:0.5) plants is higher, if we look at the mineral
content in g/100 g of dry matter (Table 5), their Ca concentration is lower. Nevertheless,
their K content is the highest of all the adult plants. This behavior is different from that
reported by Errebhi and Wilcox (1990). They found that, in all the species they tested, the
addition of NH4+ to the nutrient solution decreased the uptake of cations and increased the
uptake of anions. In our experiment, when the plant can absorb NH4+, its P uptake does not
change, and there is no general rule for the cations. However, our results agree with
Marschner (1990) that showed that K is preferentially absorbed by the plant, before Ca, and
that plant K uptake increases when Ca supply to roots decreases.
Table 5
Mineral concentrations in plants expressed as % in the whole plant dry matter and as a fraction of the N
concentration
N:K:Ca (1:1:1)
N:K:Ca (1:1:0.5)
N:K:Ca (0.5:0.5:0.5)
Ratio to N
% of DM
Ratio to N
% of DM
Ratio to N
Vegetative phase
N
1.68
P
0.41
K
3.11
Ca
1.85
Mg
0.97
1
0.2
1.9
1.1
0.6
1.54
0.40
3.09
1.51
0.90
1
0.3
2.0
1.0
0.6
1.50
0.41
3.09
1.63
0.97
1
0.3
2.1
1.1
0.6
Reproductive phase
N
1.26
P
0.30
K
1.39
Ca
2.13
Mg
1.05
1
0.2
1.1
1.7
0.8
1.94
0.27
2.48
1.84
0.86
1
0.1
1.3
0.9
0.4
1.11
0.47
1.11
2.09
1.47
1
0.4
1.0
1.9
1.3
% of DM
280
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4.4. Fertilization pattern
The importance of K supply, underlined by Boertje (1978) appears in our results. For all
the treatments, potassium is the mineral element that is absorbed in greatest amount by the
plant at every stage of its development.
In our three treatments, leaching losses exceeded supply. The nutrient use efficiency was
generally better in the reproductive phase, but the leaching losses did not decrease at all. As
shown by Cabrera (2003), significant increases in N leaching losses were observed when N
supply increased. But we did not observe the fall in fertilizer use efficiency he found with
increasing applied N. The nutrient solution that had the best nutrient use efficiency was
N:K:Ca (1:1:0.5) in the vegetative phase and N:K:Ca (1:1:1) in the reproductive phase. It
was as if the plants of the N:K:Ca (1:1:0.5) treatment could store more nutrients during the
vegetative phase.
From the plant analysis data (Table 5) of the N:K:Ca (1:1:0.5) treatment (that gave the
best yield) we propose two different nutrient solution compositions (all in mmol/l):
the first should be used during the vegetative period when the plant creates its root
system and more than a quarter of its final leaf area (Dufour and Guérin, 2003); its
composition is:
2:5 NNHþ
4 ; 5:0 NNO3 ; 1:0 P; 5:3 K; 2:6 Ca; 2:1 Mg;
the second, for the production period, contains less P and K than the first because plants
in the reproductive phase have a lower rate of P and K (Table 5):
2:5 NNHþ
4 ; 5:0 NNO3 ; 0:5 P; 3:5 K; 2:6 Ca; 2:1 Mg:
The NH4+/NO3 ratio is close to that of the N:K:Ca (1:1:0.5) solution and the total
nitrogen amount is slightly lower because nitrogen was always in excess, but it is higher
than in the solutions that gave reduced yields in Dufour and Clairon’s (1997) work, and
similar to that of Sonneveld and Voogt’s (1993) solution. On the other hand, the Sonneveld
and Voogt solution had a lower concentration of NH4+, Ca and Mg:
0:8 NNHþ
4 ; 6:6 NNO3 ; 1:0 P; 4:5 K; 1:6 Ca; 1:0 Mg:
They needed to limit NH4+ supply because of pH decrease in their substrate.
In our conditions, more than 60% of any applied nutrient was lost by leaching in the
treatment that gave the best yields. These losses should be reduced (i) by using a substrate
with a higher water holding capacity. But Anthurium andreanum Lind. is a semiepiphythic plant in its natural environment, the roots hanging in the air or growing on
moss-covered branches, so the plant is very sensitive to root rots; (ii) by decreasing the
volume of nutrient solution applied every day. But the high temperature in the greenhouse
results in a high transpiration rate and the water requirements of the plants are high.
Őzçelik and Őzkan (2002) measured a maximal water consumption of 140 ml/plant/day
for 8-month-old plants grown in Turkey. A better match between supply and demand
should be achieved with the nutrient balance method (Costa and Guérin, 1996). In this
method, the solution composition changes in relation to plant, substrate and weather
L. Dufour, V. Guérin / Scientia Horticulturae 105 (2005) 269–282
281
parameters. A more precise study could take into account an expected variation of plant
requirements connected with the seasonal variation of PAR and with leaf area expansion;
(iii) by supplying pure water between nutrient solution applications. If we consider the
plant uptake and with 30% of leaching losses, the required volume of nutrient solution per
plant is only on average 3.5 ml/day (irrigation of 0.7 103 mm) for the vegetative phase
and 40 ml/day (irrigation of 8 103 mm) for the reproductive phase. The water supply
would be far too little with these amounts; (iv) by splitting the total daily supply into
several applications.
5. Conclusion
Considering the nutrition of Anthurium andreanum, an insufficient supply of nitrogen
and potassium can severely reduce and slow down growth, increase the length of the
vegetative phase, and reduce yield. Moreover, the flowers produced are of poorer quality. A
concentration of 8.9 mmol N/l of nutrient solution is sufficient for a good yield and flower
quality. Looking to the mineral analyses, this concentration could even be reduced to
7.5 mmol/l. On the other hand, the potassium supply needs to be high, especially during the
reproductive phase, when there is intensive export from mature leaves to flowers and young
leaves due to large leaf and flower production.
Another factor affecting plant growth is the ammonium fraction of the total nitrogen
supply. This component needs further study to find the best NH4+/NO3 ratio, but our
results showed that an increase in the ammonium concentration in the nutrient solution up
to at least 1/3rd of the total nitrogen improves plant growth, development and yield.
Different fertilization schemes can be proposed in order to reduce the leaching of the
majority of the mineral elements. In particular, in our conditions, two nutrient solution
formulations can be used according to the plant’s developmental stage. Methods including
irrigation with pure water and fractionation of the supply should be tested to find the most
efficient in terms of flower yield and quality and for environmental protection.
Acknowledgements
The authors thank Dr. Jorge Sierra (Antilles-Guyane INRA), and Dr. Louis-Marie
Rivière (Angers INRA) for their very valuable advice throughout this work and for their
critical review of the manuscript. They also express their thanks to Alan Scaife for
reviewing the English.
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