Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
Pesquisas Agrárias e Ambientais
DOI: https://doi.org/10.31413/nativa.v11i2.15728
ISSN: 2318-7670
Does the film formed by the Bordeaux mixture on the leaf surface of fig trees
affect photochemical processes?
Matheus Marangon DEBASTIANI1, Angélica Alves GOMES1, Angria Ferreira DONATO1,
Mariana PIZZATTO1, Samuel Silva CARNEIRO2, Andréa Carvalho da SILVA1,2*
1 PosGraduate Program in Agronomy, Federal University of Mato Grosso, Sinop, MT, Brazil.
1 Institute of Agricultural and Environmental Sciences, Federal University of Mato Grosso, Sinop, MT, Brazil.
*E-mail: andrea.silva@ufmt.br
Submission: 06/12/2023; Accepted: 07/03/2023; Published: 07/14/2023.
ABSTRACT: This work aimed to evaluate changes in gas exchanges and chlorophyll a fluorescence in fig
plants due to the film formed on the leaf surface by Bordeaux mixture applied to control rust. The experiment
was conducted in an orchard with 7-month-old fig trees of the cultivar Roxo de Valinhos, in April 2020. A
completely randomized experimental design with four replications was used, consisting of two treatments (with
and without application of Bordeaux mixture), evaluating leaves in three different parts of the branch (apical,
middle, and basal) in five evaluations. The evaluated gas exchange parameters were: carbon, leaf temperature,
transpiration, stomatal conductance, and photosynthesis. The evaluated chlorophyll fluorescence parameters
were: maximum and effective quantum yield of the photosystem, electron transport rate, photochemical and
non-photochemical quenching, and leaf area. The film formed by Bordeaux mixture application did not affect
the photochemical phases of photosynthesis and chlorophyll a fluorescence. The leaf position on the branch
affected internal CO2 concentration and net CO2 assimilation over time. Leaves in the middle part of the
branch presented larger leaf areas than those in the apical and basal parts.
Keywords: Ficus Carica L.; chlorophyll a; rust; leaf area.
A película formada pela calda bordalesa na superfície foliar das figueiras
interfere nos processos fotoquímicos?
RESUMO: O objetivo desse trabalho foi avaliar se a camada formada pela solução da calda bordalesa na
superfície foliar altera as características das trocas gasosas, bem como a Fluorescência da clorofila A, quando
aplicada no combate a ferrugem. O delineamento experimental utilizado foi o inteiramente casualizado com 2
tratamentos (com e sem calda bordalesa) com folhas em 3 partes distintas do ramo da figueira (apical, mediana
e basal), sendo feitas 5 avaliações, com 4 repetições, em um pomar de 7 meses de idade da cultivar Roxo de
Valinhos, no mês de abril de 2020. As avaliações das trocas gasosas foram referentes a: variação do carbono,
temperatura da folha, transpiração, condutância estomática e fotossíntese. As avaliações da fluorescência da
clorofila A, foram referentes a: rendimento quântico máximo e efetivo do fotossistema, taxa de transporte de
elétrons, dissipação fotoquímica e não fotoquímica e área foliar. A película formada pela aplicação da calda
bordalesa não interferiu nas etapas fotoquímicas da fotossíntese e na fluorescência da clorofila A. A
Concentração interna, e a assimilação líquida do CO2 foram influenciadas pela posição da folha no ramo ao
longo do tempo. As folhas da parte mediana do ramo apresentam a maior área, em detrimento das partes apical
e basal.
Palavras-chave: Ficus Carica L.; clorofila A; ferrugem; área foliar.
1. INTRODUCTION
Fig trees are rustic plants that present good
edaphoclimatic adaptation, and excellent production and
vegetative performance in different regions of Brazil;
however, they are susceptible to rust disease (Cerotelium fici),
which affects their leaves and causes severe damage, resulting
in production losses and stunted plant growth (FREIRE et
al., 2006). According to Pinheiro et al. (2021) and Medeiros
(2002), it is a highly destructive and contagious disease,
whose control requires the eradication of infected leaves and
other plant parts, as well as intensive fungicide application to
affected areas for preventing infestation and plant loss;
therefore, preventive treatments and proper disposal of dead
plant material from pruning and harvest are necessary.
One of the most efficient and low-cost methods for
preventing and controlling rust on fig trees is the application
of the Bordeaux mixture. This mixture is a chemical
combination of copper sulfate and CaO, resulting in CaSO4
(calcium sulfate), which strongly adheres to leaves and has
fungicidal and bactericidal actions. Additionally, it has
nutritional functions for plants by supplying copper and
calcium through leaves, contributing to cell wall formation,
physiological defense, and enzyme synthesis (PAULUS et al.,
2001; REBELO et al., 2015).
However, the application of Bordeaux mixture leads to
the accumulation of residues on the leaf surface over time,
creating a bluish film that can thicken the leaf during dry
periods. Therefore, the objective of this study was to evaluate
Debastiani et al.
Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
201
whether the film formed on the leaf surface by the Bordeaux
mixture applied to control rust affects gas exchange and
chlorophyll a fluorescence in fig plants.
2. MATERIAL AND METHODS
This study was conducted at the plant production area of
the Federal University of Mato Grosso (UFMT), Sinop
campus, Brazil (11°51'S, 55°29' W, and altitude of 382 m), in
an orchard previously planted with fig trees of the cultivar
Roxo de Valinhos. The soil of the area was classified as Typic
Hapludox (Latossolo Vermelho-Amarelo distrofico;
SANTOS et al., 2018). The climate of the region is Aw,
tropical hot and humid, according to the Köppen
classification, with a well-defined dry season and mean
annual rainfall depth of 2,000 mm, concentrated from
October to March (SOUZA et al., 2013).
A completely randomized experimental design was used,
consisting of two treatments (with and without application of
Bordeaux mixture) to evaluate leaves in three different parts
of the branch (apical, middle, and basal) in five evaluations
(0, 1, 3, 5, and 7 days after application of Bordeaux mixture).
The experiment was conducted from April 23 to 30,
2020, in an orchard consisting of 210 seven-month-old fig
plants, spaced at 2 × 2.5 m, grown with single branches, and
irrigated through a drip system. Bordeaux mixture was
applied to four morphometrically similar plants, whereas four
other plants were grown without application. A total of 180
leaves were evaluated: 59 leaves from the upper (apical), 65
from the middle (middle), and 60 leaves from the lower
(basal) part of the branch. A Bordeaux mixture solution at
the ratio of 1:1 (copper sulfate and CaO) was prepared and
applied on April 23, 2020, following the phytosanitary
management commonly adopted by fig growers. The
Bordeaux mixture solution was applied using a pressurized
manual pump at a rate of 300 mL per plant in 30 seconds,
covering all leaves of the branch (Motta, 2008).
Gas exchanges were measured on all leaves of plants in
the treatments with and without Bordeaux mixture
application, at 0, 1, 3, 5, and 7 days after application (DAA),
using an infrared gas analyzer (IRGA) (LCi-SD; ADC
BioScientific, Hoddesdon, UK). Readings were carried out
between 8:00 am and 4:00 pm, with intervals of up to 15
minutes for cooling the device.
The leaves were placed inside the IRGA chamber,
occupying an area of 6.25 cm2, and subjected to an effective
pulse of 1839 µmol m-2 s-1 light intensity (Qleaf) to obtain the
following parameters: external carbon concentration (Cref;
μmol mol-1), internal carbon concentration (Ci; μmol mol-1),
carbon variation between the external environment and the
chamber (ΔC; μmol mol-1), leaf temperature (T; °C),
transpiration rate (E; mmol m-2 s-1), stomatal conductance (gs;
mol m-2 s-1), and net assimilation rate (A; μmol m-2 s-1).
Parameters of chlorophyll a fluorescence were evaluated
using an OS5p modulated chlorophyll fluorometer (FP-100;
Opti-Sciences, Hudson, USA), according to the following
analysis protocols: maximum quantum yield (Fv/Fm
protocol; measured in the dark-adapted state; and effective
quantum yield (yield protocol), measured in the light-adapted
state. The measurements were carried out at 0, 1, 3, and 7
DAA on a smaller number of leaves (45 apical leaves, 48
middle leaves, and 45 basal leaves) to obtain the following
parameters of chlorophyll a fluorescence: effective quantum
yield of PSII (ΦPSII), electron transport rate (ETR),
photochemical quenching (qP), non-photochemical
quenching (NPQ), and maximum quantum yield of PSII
(Fv/Fm).
Leaf area (LA) was assessed at the first and last
evaluations by measuring length and width (cm) and then
applying the equation proposed by Souza et al. (2014).
The data were subjected to the Shapiro-Wilk normality
test, analysis of variance at 1% and 5% probability levels, and
Scott-Knott test, using the software SISVAR.
The climate data were obtained from a weather station
installed in the UFMT, Sinop. The air temperature remained
within the range of 20.5 to 33.5 °C, with means varying from
24 to 26 °C, within the optimal range for crop development:
20 to 25 °C (SOUZA and LEONEL, 2011). The relative air
humidity decreased from the first (0 DAA) to the last (7
DAA) evaluation. The daily curves of photosynthetically
active radiation (PAR), global radiation, and illuminance
(Lux) exhibited the expected dynamics, with higher
intensities between 10:00 am and 1:00 pm.
3. RESULTS
The analysis of variance (Table 1) showed a significant
triple interaction (treatments, leaf position on the branch, and
evaluation day) for stomatal conductance (gs) (gas exchange
parameter) and quantum yield of PSII and electron transport
rate (chlorophyll a fluorescence parameters).
Gas exchange parameters were not significantly affected
by the Bordeaux mixture application; significant variations
were found only for leaf position and evaluation day.
Therefore, the comparison between treatments with and
without Bordeaux mixture application was not necessary
throughout the discussion.
Leaf internal temperature presented no statistically
significant variation for treatments and branch parts (Table
2). Variations in CO2 concentration between leaf external and
internal environments (ΔC) and net CO2 assimilation rate (A)
presented higher means for leaves in the apical part of the
branch (29.48 to 42.33 μmol mol-1 and 9.75 to 13.68 μmol m-
2s-1, respectively), followed by leaves in the middle part (25.59
to 33.84 μmol mol-1 and 8.62 to 11.06 μmol m-2s-1) and basal
part of the branch (19.08 to 26.87 μmol mol-1 and 6.34 to
8.74 μmol m-2s-1, respectively). Transpiration rate and
stomatal conductance showed low variation for leaf positions
on the branch.
Significant statistical variation was found for gs (Table 1)
for treatments with and without Bordeaux mixture
application, mainly in apical leaves, except for the second
evaluation (Table 2).
Chlorophyll A fluorescence showed no significant triple
interaction for maximum photochemical efficiency of PSII
(Fv/Fm), photochemical quenching (qP), and non-
photochemical quenching (NPQ); however, significant
statistical variation was found for quantum yield of PSII
(ΦPSII) and electron transport rate (ETR).
Chlorophyll a fluorescence parameters were not
significantly affected by Bordeaux mixture application; the
treatments and evaluation day were the factors with higher
effects; therefore, discussions for treatments were
unnecessary since no significant variation was found.
Does the film formed by the Bordeaux mixture on the leaf surface of fig trees affects photochemical ...
Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
202
Table 1. Analysis of variance for parameters of gas exchange and chlorophyll a fluorescence: external carbon concentration (Cref), internal carbon
concentration (Ci), carbon variation between the external environment and the analysis chamber (ΔC), leaf temperature (T), transpiration rate (E),
stomatal conductance (gs), net assimilation rate (A), maximum photochemical efficiency of PSII (Fv/Fm), photochemical quenching (qP
), non-
photochemical quenching (NPQ), effective quantum yield of PSII (ΦPSII), and electron transport rate (ETR). Leaf area in the treatments with
Bordeaux mixture application (BM) and without Bordeaux mixture application (WBM) in fig trees (cultivar Roxo de Valinhos).
Tabela 1. Análise de variância para os parâmetros das trocas gasosas e fluorescência da clorofila A referentes a: concentração de carbono no ambiente
externo (Cref), concentração de carbono interno (Ci), variação de carbono entre ambiente externo e câmara de análise (ΔC), temperatura da folha
(TF), taxa de transpiração (E), condutância estomática (gs) e taxa de assimilação líquida (A); máxima eficiência fotoquímica de PSII (Fv/Fm),
Quenching fotoquímico (qP
), Dissipação não fotoquímica (NPQ), Rendimento quântico efetivo do PSII (Φ PSII) e Taxa de transporte de elétrons
(ETR). Área foliar nos tratamentos com aplicação de calda bordalesa (CC) e sem aplicação de calda bordalesa (SC), em plantas de Figueiras ‘Roxo
de Valinhos’.
parameters of gas exchanges
Source of variation
Cref
Δ
C
i
T
E
g
s
A
Treatment (Treat)
5.136**
49.884*
1.269
NS
1.807
NS
5.301**
21.497*
46.928*
Leaf position (Leaf) 2.919NS 171.550* 126.701* 2.708NS 26.089* 17.146* 166.555*
Evaluation day (Day) 64.094* 11.037* 17.565* 9.717* 27.188* 25.959* 10.207*
Treat×Leaf 0.877NS 2.971NS 0.328NS 0.949NS 2.976NS
11.734* 2.704NS
Treat×Day 3.728* 1.136NS 7.399* 1.264NS 3.507* 6.189* 1.177NS
Part×Day 0.307NS 1.344NS 1.091NS 1.415NS 1.025NS
1.356NS 1.285NS
Treat×Leaf×Day 0.362NS 1.242NS 1.423NS 1.257NS 0.419NS
2.169** 1.184NS
Parameters of Chlorophyll
a
Fluorescence
Leaf Area
Source of variation
Fv/Fm
q
P
NPQ
ΦPSII
ETR
BM
WBM
Treatment (Treat)
16.366*
0.848
NS
0.086
NS
2.270
NS
0.145
NS
-
-
Leaf position (Leaf) 15.550* 8.781** 1.941NS 6.505* 4.127* 58.589** 90.892**
Evaluation day (Day) 2.978** 4.167** 13.430** 2.319NS 3.740* - -
Treat×Leaf 1.798NS 1.299NS 2.694NS 1.478NS 0.843NS
2.547NS 0.874NS
Treat×Day 0.559NS 1.143NS 1.459NS 1.375NS 1.441NS
- -
Part×Day 1.221NS 4.798** 0.779NS 4.698* 2.271* 3.666* 3.995*
Treat×Leaf×Day 1.249NS 2.070NS 0.076NS 3.017* 2.474*
* = significant at 5%; ** = significant at 1%; NS = not significant.
Table 2. Parameters of gas exchanges as a function of days after application (DAA) of Bordeaux mixture in fig trees (cultivar Roxo de Valinhos):
external carbon concentration (Cref), internal carbon concentration (Ci), carbon variation between the external environment and the analysis
chamber (ΔC), leaf temperature (T), transpiration rate (E), stomatal conductance (gs), and net assimilation rate (A).
Tabela 2. Parâmetros de troca gasosas em função dos dias após aplicação ou não da calda bordalesa, em plantas de Figueiras ‘Roxo de Valinhos’:
concentração de carbono no ambiente externo (Cref), concentração de carbono interno (Ci), variação de carbono entre ambiente externo e câmara
de análise (ΔC), temperatura da folha (TF), taxa de transpiração (E), condutância estomática (gs) e taxa de assimilação líquida (A).
Leaf position on
DAA
Cref
C
i
C
T
E
g
s
A
Bordeaux Mixture Application
Apical
0
378.07 Ba
283.24 Bc*
29.48 Aa
37.49 Aa*
4.69 Bb
0.29 Cc*
9.75 Aa
1
378.26 Ba
306.17 Ab*
31.82 Aa*
34.10 Bb*
4.70 Ba*
0.53 Aa
10.37 Aa*
3
381.63 Ba
298.39 Ac
32.05 Aa
35.82 Ba
4.73 Bb*
0.41 Bb*
10.45 Aa
5
387.01 Aa
286.20 Bb
34.86 Aa*
37.49 Aa
5.71 Ab
0.35 Bb*
11.38 Aa*
7
392.11 Aa
302.30 Ab
32.72 Aa*
35.10 Ba
4.94 Bb
0.37 Bb*
10.72 Aa*
Middle
0
377.87 Ba
302.01 Bb
26.82 Aa
36.88 Aa
5.20 Ba
0.37 Bb*
8.78 Aa
1
375.80 Ba
313.21 Ab*
26.10 Ab*
37.95 Aa*
4.91 Ba*
0.49 Aa
8.62 Ab*
3
377.97 Ba
314.03 Ab
27.11 Ab
35.76 Ba
5.09 Bb
0.53 Aa
8.97 Ab
5
384.37 Aa*
292.70 Bb
30.09 Ab
38.17 Aa
5.91 Ab
0.32 Bb
9.82 Ab
7
387.60 Aa
324.53 Aa
25.59 Ab
35.27 Ba
5.28 Ba
0.50 Aa
8.38 Ab*
Basal
0
377.72 Ba
324.66 Aa
19.08 Ab*
36.83 Aa
5.54 Ba
0.45 Ba
6.34 Ab*
1
377.32 Ba
326.88 Aa
22.04 Ac
35.25 Ab
5.10 Ba
0.56 Aa
7.23 Ac
3
376.28 Ba
327.09 Aa
20.71 Ac
36.34 Aa
5.66 Ba
0.57 Aa
6.82 Ac
5
385.37 Aa
315.73 Ba
24.59 Ac
37.47 Aa
6.32 Aa
0.44 Ba
8.13 Ac
7
386.26 Aa
335.13 Aa
19.49 Ac
35.60 Aa
5.53 Ba
0.50 Ba
6.56 Ac
Without Bordeaux Mixture Application
Apical
0
380.23 Ba
299.76 Ab*
32.48 Ba
35.46 Aa*
4.79 Ba
0.53 Aa*
10.61 Ba
1
374.54 Ca
287.29 Bb*
37.94 Aa*
35.90 Aa*
5.37 Ba*
0.49 Aa
12.34 Aa*
3
381.11 Ba
299.82 Ab
35.80 Ba
36.08 Aa
5.21 Ba*
0.52 Aa*
11.67 Ba
5
390.55 Aa
287.45 Bc
42.33 Aa*
36.58 Aa
5.92 Aa
0.46 Aa*
13.68 Aa*
7
391.86 Aa
299.47 Ac
40.11 Aa*
34.54 Aa
5.19 Bb
0.51 Aa*
13.01 Aa*
Middle
0
378.54 Ba
308.30 Ab
27.79 Bb
35.98 Aa
5.10 Ba
0.47 Aa*
9.25 Bb
1
373.60 Ca
293.80 Bb*
33.84 Aa*
36.21 Aa*
5.45 Aa*
0.49 Aa
11.06 Aa*
3
380.48 Ba
316.88 Aa
27.17 Bb
35.87 Aa
4.90 Ba
0.52 Aa
8.77 Bb
5
392.30 Aa*
303.82 Bb
32.22 Ab
37.00 Aa
5.75 Aa
0.37 Ba
10.52 Ab
7
392.08 Aa
315.39 Ab
29.54 Bb
35.55 Aa
5.37 Ab
0.43 Ba
9.73 Bb*
Basal
0
377.86 Ba
319.62 Ba
23.90 Ab*
36.52 Aa
5.37 Ba
0.51 Aa
7.86 Ac*
1
374.35 Ba
316.95 Aa
23.76 Ab
35.76 Aa
5.50 Ba
0.56 Aa
7.88 Ab
3
378.90 Ba
327.12 Ba
22.22 Ac
36.33 Aa
5.42 Ba
0.58 Aa
7.35 Ac
5
389.82 Aa
317.41 Ba
26.87 Ac
37.42 Aa
6.18 Aa
0.41 Ba
8.74 Ac
7
391.29 Aa
333.51 Aa
23.69 Ac
35.70 Aa
5.87 Aa
0.51 Aa
7.76 Ac
*Means followed by the same letter in the row and in the column not statistically different by test the Skott-Knott test at 5% probability level.
Debastiani et al.
Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
203
Fv/Fm presented sporadic variations for evaluation days
(Table 3), with decreases in apical and middle leaves on
isolate days, denoting uniformity of results throughout the
evaluations. The parameters qP, Fv/Fm, NPQ presented
significant statistical differences at 7 DAA for the treatments.
Basal leaves in both treatments (with and without Bordeaux
mixture application) presented lower qP means (0.063 and
0.054, respectively) compared to middle (0.087 and 0.126,
respectively) and apical (0.265 and 0.164, respectively) leaves
(Table 3).
A similar result was found for NPQ, which showed
significant variation only between evaluations of each
treatment, mainly for the treatment with Bordeaux mixture
application, which presented the highest mean at 0 DAA,
whereas the other evaluations presented no significant
differences from each other.
The triple interaction was significant for ETR and ΦPSII
at 1% and 5% significance levels, respectively, as both
parameters are closely correlated. ETR considers ΦPSII in its
base equation, which provides consistent results.
The absence of significant statistical variations in the
other evaluations of ETR and ΦPSII was similar for qP,
NPQ, and Fv/Fm, which showed normality, with isolated
significant statistical variations that did not unfold over time
for leaf positions and treatments.
Leaf area (LA) showed significant statistical variation for
evaluation days (0 DAA and 7 DAA) and leaf positions on
the branch (Table 1). Regarding variations in LA for
evaluation days, middle leaves in the treatment with
Bordeaux mixture application showed decreases in LA from
the first to the second evaluation, whereas apical leaves in the
treatment without Bordeaux mixture application showed
increases in LA from the first to the second evaluation.
Regarding variations in LA for leaf positions on the branch,
LA varied in both evaluations, with middle leaves showing
the highest LA means (Table 4).
Table 3. Means of maximum photochemical efficiency of PSII (Fv/Fm), photochemical quenching (qP), non-photochemical quenching (NPQ),
effective quantum yield of PSII (ΦPSII), and electron transport rate (ETR) referring to treatments (with and without Bordeaux mixture application),
evaluation days (days after application – DAA), and leaf position on the branch (apical, middle, and basal parts) of fig trees.
Tabela 3. Valores médios da Máxima eficiência fotoquímica de PSII (Fv/Fm); Quenching fotoquímico (qP); Dissipação não fotoquímica (NPQ);
Rendimento quântico efetivo do PSII (Φ PSII) e Taxa de transporte de elétrons (ETR), referentes aos tratamentos, dias e segmentos da figueira.
Leaf position on
the branch
DAA
Fv/Fm
qP
NPQ
ΦPSII
ETR
With Bordeaux Mixture Application
Apical
0
0.756 Aa
0.138 Ba
1.027 Aa
0.077 Ba
19.58 Aa
1
0.755 Aa*
0.105 Ba
0.523 Ba*
0.070 Ba
17.800 Aa
3
0.742 Ab
0.107 Ba
0.766 Ba
0.060 Ba
15.275 Aa
7
0.717 Ab
0.265 Aa*
0.595 Ba
0.170 Aa*
20.120 Aa
Middle
0
0.746 Aa*
0.163 Aa
1.208 Aa
0.086 Aa
21.668 Aa
1
0.751 Aa
0.108 Aa
0.744 Ba
0.066 Aa
16.604 Aa
3
0.749 Ab
0.089 Aa
0.790 Ba
0.052 Aa
13.204 Aa
7
0.718 Ab
0.087 Ab
0.837 Ba
0.053 Ab
13.413 Ab
Basal
0
0.752 Aa
0.153 Aa*
1.230 Aa
0.085 Aa*
21.515 Aa*
1
0.761 Aa
0.077 Aa
0.594 Ba
0.051 Aa
12.840 Ab
3
0.788 Aa
0.112 Aa
0.843 Ba
0.072 Aa
18.181 Aa
7
0.757 Aa
0.063 Ab
0.542 Ba
0.040 Ab
10.030 Bb
Without bordeaux
mixture application
Apical
0
0.730 Aa
0.167 Aa
1.036 Aa
0.097 Aa
24.591 Aa
1
0.689 Ab*
0.105 Ba
0.867 Aa*
0.055 Aa
13.952 Ba
3
0.713 Ab
0.084 Ba
0.904 Aa
0.048 Aa
12.177 Ba
7
0.698 Aa
0.164 Aa*
0.785 Aa
0.082 Aa*
20.855 Aa
Middle
0
0.763 Ab*
0.142 Aa
1.049 Aa
0.065 Ab
16.386 Ab
1
0.753 Aa
0.136 Aa
0.786 Aa
0.080 Aa
20.176 Aa
3
0.765 Ab
0.100 Aa
0.698 Aa
0.059 Aa
14.876 Aa
7
0.745 Aa
0.126 Aa
0.857 Aa
0.070 Aa
17.659 Aa
Basal
0
0.695 Aa
0.065 Ab
0.919 Aa
0.039 Ab*
9.977 Bb*
1
0.732 Aa
0.112 Aa
0.519 Aa
0.079 Aa
19.963 Aa
3
0.732 Aa
0.079 Aa
0.787 Aa
0.050 Aa
12.594 Ba
7
0.713 Aa
0.054 Ab
0.634 Aa
0.034 Aa
8.561 Bb
*Means followed by the same letter in the row and in the column are not statistically different by the Skott-Knott test at 5% probability level.
Table 4. Means of leaf area (cm²) in treatments with and without Bordeaux mixture application in fig trees (cultivar Roxo de Valinhos).
Tabela 4. Área foliar média dos tratamentos com aplicação de calda bordalesa e sem aplicação de calda bordalesa, em plantas de Figueiras ‘Roxo de
Valinhos’.
DAA With Bordeaux Mixture Application Without Bordeaux Mixture Application
Apical leaves Middle leaves Basal leaves Apical leaves Middle leaves Basal leaves
0 270.04 Ba 359.73 Aa 200.75 Ca 309.02 Bb 377.82 Aa 195.25 Ca
7 294.97 Aa 309.20 Ab 183.31 Ba 367.15 Aa 362.47 Aa 186.72 Ba
* Means followed by the same letter in the row and column are not statistically different by the Skott-Knott test at 5% probability level.
4. DISCUSSION
Internal leaf temperature remained approximately 2 to 3
°C higher than the maximum environmental temperatures
during the experiment (Figure 1). According to Taiz et al.
(2017), this is due to continuous exposure to solar radiation
and bright environmental conditions; this excessive
accumulation is undesirable for plants, as high temperatures
affect water balance and CO2 assimilation.
Does the film formed by the Bordeaux mixture on the leaf surface of fig trees affects photochemical ...
Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
204
Internal CO2 concentration is a limiting factor for
photosynthesis and stomatal regulation, mainly in C3
photosynthetic metabolism plants such as fig trees. Once
inside the leaf, CO2 diffuses from the intercellular airspaces
to the chloroplast and is limited by resistances in gas and
liquid phases of cytosol, as well as diffusion barriers, causing
it to accumulate in intercellular spaces, mainly when stomatal
conductance and assimilation rate are affected by external
factors, such as air temperature and humidity (CHAVES et
al., 2011; TAIZ et al., 2017).
Silva et al. (2010) evaluated gas exchange in young leaves
(recently opened and near the branch meristem) of fig trees
and found that the lowest net CO2 assimilation rates,
transpiration, and stomatal conductance resulted in high CO2
concentration in the substomatal chamber and low
photosynthetic carbon assimilation. However, the results
found in the present study showed that apical leaves had the
lowest internal CO2 concentrations and the highest means of
net CO2 assimilation, i.e., carbon dioxide was moving
towards the carboxylation stage of the photosynthetic
process, followed by middle and basal leaves.
Transpiration rate (E) refers to water loss through
stomatal pores, combined with the guard cells, at the time of
opening. González-Rodríguez and Peters (2010) evaluated
leaf sprouting in pruned and unpruned fig trees in Spain and
found transpiration rates ranging from 3 to 7.6 mmol m-2 s-1
over 200 days; the highest rates were found during the
summer and were similar to the range of 4.6 to 6.1 mmol m-
2 s-1 found in the present study (Table 2). Additionally, Silva
et al. (2010), assessed gas exchange in leaves of fig trees of
the cultivar the Roxo de Valinhos in Botucatu, SP, Brazil, and
found transpiration rates ranging from 2.19 to 4.78 mmol m-
2 s-1, indicating that the transpiration rates found in the
present study are consistent with those previously reported
for the species.
The means of stomatal conductance (gs) varied from
0.280 to 0.584 mol m-2 s-1, which are similar to results
reported in the literature. Ammar et al. (2020) evaluated
physiological dynamics in 5-year-old fig trees over 259 days
in Tunisia and found maximum means of 0.370 and 0.435
mol m-2 s-1 during late spring when temperatures ranged from
28 to 31 °C; however, the lowest means (below 0.100 mol m-
2 s-1) were found during summer when temperatures were
higher than 35 °C.
Can et al. (2008) and Ammar et al. (2020) reported that
the maximum gs was found when daytime temperatures were
around 30 to 32 °C, as at these temperatures, water viscosity
decreases and mesophyll conductance increases, increasing
guard cell turgor pressure and stomatal opening. Conversely,
gas exchange rates were significantly lower during the hottest
periods of the year compared to those found in early summer.
The net assimilation rate showed localized variations in
leaf positions on the branch regarding the evaluation days,
specifically in apical and middle leaves in the treatment
without Bordeaux mixture application at 0 and 3 DAA.
Regarding the variation among leaf positions within each
treatment, a significant difference was found, with the highest
rates in apical leaves and the lowest rates in basal leaves were
inversely proportional to the results found for internal
carbon concentration (Ci).
The results obtained differed from those reported by
Silva et al. (2010), who found the lowest rates of net CO2,
transpiration, and stomatal conductance in younger leaves,
resulting in higher CO2 accumulation in the substomatal
chamber. Only stomatal conductance and transpiration rate
remained within the expected range, with the lowest rates
found in apical leaves in the present study.
The lower CO2 concentration found in apical leaves,
combined with the higher assimilation rate, may be related to
environmental conditions, mainly intense light and radiation,
as leaf temperatures did not vary significantly and the lowest
transpiration rate and stomatal conductance were found in
apical leaves, denoting that the leaves maintained the CO2
assimilation even with partially open stomata.
The mean assimilation rates found (6.345 to 13.018 μmol
m-2 s-1) are consistent with results reported in the literature
for fig tree crops. Costa et al. (2020) found rates ranging from
6.06 to 10.49 μmol m-2 s-1 when studying photosynthetic
dynamics in different numbers of branches of fig trees
(cultivar Roxo de Valinhos) in Erechim, RS, Brazil.
According to Ferraz et al. (2020), the net assimilation rate
represents the photosynthesis performed by the plant.
Therefore, superior performance in CO2 assimilation results
in increased quantum efficiency and better utilization and
conversion into light energy, leading to higher allocation of
biomass and the formation of better plant architecture. The
authors compared commercial accessions of fig trees and
found means ranging from 5.74 to 12.94 μmol m-2 s-1 and a
mean of approximately 12.59 μmol m-2 s-1 for the cultivar
Roxo de Valinhos, which is similar to that found in the
present study.
Regarding the evaluations of chlorophyll a fluorescence,
the maximum photochemical efficiency of PSII (Fv/Fm)
varied over time in the treatments, mainly with decreases in
Fv/Fm on isolate evaluation days in the apical and middle
leaves, without significant variations. These results were
expected since characteristics such as cultivar, time, and leaf
position on the branch do not affect Fv/Fm, but significant
variations are due to environmental factors such as high
radiation incidence (PALLIOTTI, et al. 2009).
The correlation between maximum fluorescence (Fm)
and variable fluorescence (Fv) enables a better understanding
of qualitative and quantitative analyses of light energy
absorption and use by the photosystem II (PS II); it is an
indicator of use efficiency of photochemical radiation and,
consequently, carbon assimilation by plants, contributing to
the diagnosis of integrity of the photosynthetic apparatus
after exposure to environmental adversities, especially when
combined with gas exchange analyses (TATAGIBA et al.,
2014; FREIRE et al., 2014).
The means found for Fv/Fm (0.689 to 0.788) were
consistent with other Fv/Fm results reported for arboreal
species. Ammar et al. (2020) found means ranging from 0.493
to 0.741 in an experiment in Tunisia, with the lowest means
found in summer and early autumn when temperatures and
light incidence were higher. Similarly, Gomes et al. (2008)
found Fv/Fm means close to 0.741 in a study characterizing
the photosynthetic performance of fig trees in the state of
Espirito Santo, Brazil; the authors noted that other fruit tree
species grown in the same area, such as coconut and mango
trees, had Fv/Fm means of 0.74 and 0.76, respectively, which
are similar to the results found in the present study.
The triple interaction was significant for the electron
transport rate (ETR) and effective quantum yield of PSII
(ΦPSII) at 1% and 5% probability levels, respectively. It was
significant for ΦPSII due to apical leaves, as the treatment
Debastiani et al.
Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
205
with Bordeaux mixture application was the only one that
showed variation among evaluations for apical leaves. The
highest mean (0.170) was found at 7 DAA. Apical leaves had
higher ΦPSII at 7 DAA than middle and basal leaves, which
did not differ from each other (0.053 and 0.040, respectively).
Apical and basal leaves presented significant variation
between treatments at 0 DAA; apical leaves in the treatment
with Bordeaux mixture application had higher ΦPSI at 7
DAA than those in the treatment without application, which
did not differ among evaluations and leaf positions on the
branch.
Similar results were found for ETR in basal leaves in the
treatment without Bordeaux mixture application. Regarding
the evaluation days within each treatment, the lowest ETR
means were found at the 0 and 7 DAA. Significant variation
between treatments was found only for basal leaves in the
first evaluation, with the highest mean (21.52 μmol m-2 s-1)
found in the treatment with Bordeaux mixture application.
The absence of significant statistical variations in ETR
and ΦPSII in the other evaluations is consistent with the
results of qP, NPQ, and Fv/Fm, which showed normality
with isolated significant variations that did not unfold over
time within leaf positions and treatments.
The results found for all parameters, except Fv/Fm, were
lower compared to those reported by Ranjbar-Fordoei
(2019), who evaluated three stages of fig leaf ontogeny
(young, mature, and senescent) in Iran for two production
years and found qP and ΦPSII with significant differences in
leaf growth and maturation, which was not found in the
present study, and higher mean for all evaluations of qP
(0.370 to 0.544) and ΦPSII (0.454 0.502); senescent leaves
presented the lowest values.
According to Moreno et al. (2008) and Ranjbar-Fordoei
(2018), low qP and ΦPSII (and consequently ETR) are
associated with abiotic stresses, such as water stress,
indicating that the PSII photosynthetic apparatus may have
been damaged and lost its ability to dissipate heat, resulting
in low efficiency in light energy transformation in PSII,
mainly in the primary light capture when PSII reaction
centers are partially deactivated; similarly, a decrease in qP
may indicate damage to PSII reaction centers and collapse in
the balance between excitation rate and electron transfer rate.
Leaf area (LA) differed significantly among leaf positions
on the branch in both evaluations. Middle leaves had the
highest mean LA, followed by apical and basal leaves. In the
evaluation at 7 DAA, apical and middle leaves did not show
significant differences from each other for LA.
Ferraz et al. (2020) reported that fig trees maintain
constant leaf gains and LA until harvest and dormancy of the
plant when these values decrease.
5. CONCLUSIONS
The film formed on the leaf surface due to the application
of Bordeaux mixture did not affect photochemical stages of
photosynthesis and chlorophyll a fluorescence.
Leaf temperature, transpiration, and stomatal
conductance presented no significant variations in gas
exchange among leaf positions on the branch (apical, middle,
and basal parts) within each treatment.
Internal CO2 concentration and net CO2 assimilation
were gas exchange parameters affected by the leaf position
on the branch over time.
Leaves in the middle part of the branch had larger leaf
areas compared to those in the apical and basal parts of the
branch.
6. REFERÊNCIAS
AMMAR, A; AISSA. I. B.; MARS, M.; GOUIAA, M.
Seasonal variation of fig tree (Ficus carica L.) physiological
characteristics reveals its adaptation performance. South
African Journal of Botany, v. 132, p 30-37, 2020.
https://doi.org/10.1016/j.sajb.2020.04.020.
CAN, H. Z.; MEYVACI, K. B.; BALCI, B. Determination of
gas exchange capacity of some Breba Fig Cultivars. Acta
Horticulturae, v. 798, p. 117-122, 2008.
https://doi.org/10.17660/ActaHortic.2008.798.14.
CHAVES, M. M.; COSTA, M.; SAIBO, N. J. M. Recent
advances in photosynthesis under drought and salinity.
Advances in Botanical Research, v. 57, p. 49-104,
2011. https://doi.org/10.1016/B978-0-12-387692-
8.00003-5
COSTA, T.; GIACOBBO, C. L.; GALON, L.; FORTE, C.
T.; DAMIS, R.; TIRONI, S. P. Management of soil cover
and its influence on phytosociology, physiology and fig
production. Comunicata Scientiae, v. 11, p. 1-10, 2020.
https://doi.org/10.14295/cs.v11i0.3236
FERRAZ, R. A.; LEONEL, S.; SOUZA, J. M. A.;
FERREIRA. R. B.; MODESTO, J. H.; ARRUDA, L. L.
Phenology, vegetative growth, and yield performance of
fig in Southeastern Brazil. Pesquisa Agropecuária
Brasileira, v. 55, e01192, 2020.
https://doi.org/10.1590/S1678-
3921.pab2020.v55.01192.
FREIRE, F. C. O.; PARENTE, G. B.; CARDOSO, B. B.
Doenças da Figueira (
Ficus carica
L.) no Estado do
Ceará. Fortaleza: Embrapa Agricultura Tropical, 2006.
5p. (Circular Técnica On-Line, 26). Available on:
https://ainfo.cnptia.embrapa.br/digital/bitstream/CNP
AT-2010/11980/1/Ci-026.pdf
FREIRE, J. L. O.; DIAS, T. J.; CAVALCANTE, L. F.;
FERNANDES, P. D.; NETO, A. J. L. Rendimento
quântico e trocas gasosas em maracujazeiro amarelo sob
salinidade hídrica, biofertilização e cobertura morta,
Revista Ciência Agronômica, v. 45, n. 1, p. 82-91, 2014.
https://doi.org/10.1590/S1806-66902014000100011
GOMES, T. D. U. H.; MENGARDA, L. H. G.; COSTA, A.
N. da; CAETANO, L. C. S.; COSTA, A. de F. S. da.;
SILVA, D. M. Caracterização fotossintética de figo (Ficus
carica L.). In: CONGRESSO BRASILEIRO DE
FRUTICULTURA, XX; Annual Meeting Of The
Interamerican Society For Tropical Horticulture, LIV.
Anais... Vitória: INCAPER: Sociedade Brasileira de
Fruticultura, 2008. 5p. Available on:
https://biblioteca.incaper.es.gov.br/digital/bitstream/it
em/152/1/CARACTERIZACAO-
FOTOSSINTETICA-DE-FIGO-Ficus-carica-L-CD-
ANAISsmallpdf.com.pdf
GONZÁLEZ-RODRÍGUEZ, A. M.; PETERS, J. Strategies
of leaf expansion in Ficus carica under semiarid
conditions. Plant Biology, v. 12, n. 3, p. 469-274, 2010.
https://doi.org/10.1111/j.1438-8677.2009.00220.x
MEDEIROS, A. R. M. de. Figueira (
Ficus carica
L.) do
plantio ao processamento caseiro. Pelotas:
EMBRAPA Clima Temperado, 2002. 16p. (Circular
Técnica On-Line, 35). Available on:
Does the film formed by the Bordeaux mixture on the leaf surface of fig trees affects photochemical ...
Nativa, Sinop, v. 11, n. 2, p. 200-206, 2023.
206
https://www.infoteca.cnptia.embrapa.br/infoteca/bitstr
eam/doc/743511/1/circular35.pdf
MORENO, S. G.; VELA, H. P.; ALVAREZ, M. O. S. La
fluorescencia de la clorofila a como herramienta en la
investigación de efectos tóxicos en el aparato
fotosintético de plantas y algas. Revista de Educación
Bioquímica, v. 27, n. 4, p. 119-129, 2008.
MOTTA, I. S. Calda bordalesa: utilidades e preparo.
Dourados: Embrapa Agropecuária Oeste, 2008. 2p.
Available on:
https://ainfo.cnptia.embrapa.br/digital/bitstream/item
/38833/1/FOL200837.pdf
PALLIOTTI, A.; SILVESTRONI. O.; PETOUMENOU,
D. Photosynthetic and photoinhibition behavior of two
field grown grapevine cultivars under multiple summer
stresses. American Journal of Enology and
Viticulture, v. 60, n. 2, p. 189-198, 2009.
https://doi.org/10.5344/ajev.2009.60.2.189
PAULUS, G.; MULLER, A. M.; BARCELOS, L. A. R;
Preparo e uso da calda bordalesa. Agroecologia e
Desenvolvimento rural Sustentável, v. 2, n. 2, p. 01-02,
2001. (Coordenação técnica: EMATER/RS, Coleção
Aprendendo a Fazer Melhor). Available on:
https://www.projetovidanocampo.com.br/agroecologia
/preparo_da_calda_bordalesa.pdf
PINHEIRO, S. O.; MELO, B.; MANCIN, C. A. Cultura da
Figueira. Uberlândia: Instituto de Ciências Agrárias
(ICIAG), 2021. (Website) Disponível em:
http://www.fruticultura.iciag.ufu.br/figo. Acesso em: 28
de setembro de 2021.
RANJBAR-FORDOEI, A. Comparative functioning of
photosynthetic apparatus and leaf water potential in
(Zygophyllum eurypterum) during phenological phases and
summer drought. Desert Ecosystem Engineering
Journal, v. 1, n. 1, p. 53-60, 2018. DOI:
https://doi.org/10.22052/jdee.2017.63258.
RANJBAR-FORDOEI, A. Impacts of elevational changes
and leaf maturity stages on photoprotective strategies and
biochemical traits of wild fig [Ficus Carica Subsp.
Rupestris (Hausskn)]. International Journal of Fruit
Science, v. 20, n. 4, p. 768-785, 2019.
https://doi.org/10.1080/15538362.2019.1673874
REBELO, J. A.; REBELO, A. M.; SCHALLENBERGER,
E. Calda bordalesa: componentes, obtenção e
características. Florianópolis: Epagri, 2015. 36p.
(Epagri. Boletim Técnico, 166). Available on:
https://publicacoes.epagri.sc.gov.br/BT/article/view/4
16
SANTOS, H. G. dos; JACOMINE, P. K. T; ANJOS, L. H.
C. dos; OLIVEIRA, V. A. de; LUMBRERAS, J. F.;
COELHO, M. R.; ALMEIDA, J. A. de; ARAUJO
FILHO, J. C. de; OLIVEIRA, J. B. de; CUNHA, T. J. F.
Brazilian Soil Classification System. 5th ed. Brasília,
DF: Embrapa, 2018.
SILVA, A. C.; LEONEL S.; SOUZA, A. P.; DOMINGOS,
J. R.; DUCATTI, C. Trocas gasosas e ciclo fotossintético
da figueira “Roxo de Valinhos.” Ciência Rural, v. 40, n.
6, p. 1270-1276, 2010. https://doi.org/10.1590/S0103-
84782010000600005
SOUZA, A. P.; MOTA, L. L.; ZAMADEI, T.; MARTIN, C.
C.; ALMEIDA, F. T.; PAULINO, J. Classificação
climática e balanço hídrico climatológico no estado de
Mato Grosso. Nativa, 1:34-43, 2013.
http://doi.org/10.31413/nativa.v1i1.1334
SOUZA, A. P.; SILVA, A. C.; LEONEL, S.; SOUZA, M. E.;
TANAKA, A. A. Estimativas da área da folha de figueiras
‘Roxo de Valinhos’ usando dimensões lineares do limbo
foliar. Ciência Rural, v. 44, n. 7, p. 1172-1179, 2014.
http://dx.doi.org/10.1590/0103-8478cr20130699.
SOUZA, M. E.; LEONEL. S. Propagação da figueira. In:
LEONEL. S.; SAMPAIO. A. C. A figueira. São Paulo:
Editora Unesp, 2011. p. 77-92.
TAIZ, L.; ZEIGER, E.; MOLLER, I.; MURPHY, A.
Fisiologia e desenvolvimento vegetal. 6 ed. Porto
Alegre: Artmed, 2017. 888p.
TATAGIBA, S. D.; MORAES, G. A. B. K.;
NASCIMENTO, K. J. T.; PELOSO, A. F. Limitações
fotossintéticas em folhas de plantas de tomateiro
submetidas a crescentes concentrações salinas,
Engenharia na agricultura, v. 22, n. 2, p. 138-149, 2014.
https://doi.org/10.13083/1414-3984.v22n02a05
Acknowledgments: The authors thank the Brazilian Coordination
for the Improvement of Higher Education Personnel (CAPES) for
granting scholarship during the conduction of the experiment; the
graduate program in Agronomy; and the members of the research
group "Environment and Plant Interaction".
Author Contributions: M.M.D. field data collection, laboratory
analysis, statistical analysis, initial writing; A.A.G.; A.F.D.; M.P. and
S.S.C. field data collection; A.C.S. - conceptualization,
methodology, research, validation, writing draft. All authors read
and agreed to the published version of the manuscript.
Funding: PROPEQ/UFMT 2021 (Researcher Assistance
Announcement – Process: 23108089302/2021-59).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Raw and analyzed data can be
obtained by request to the corresponding Author by e-mail.
Conflicts of Interest: The authors declare that there is no conflict
of interest regarding the publication of this paper.