Ojip

Derivate OJIP Variables in Sugarcane to Predict Cane Weight, Sucrose
Content, and Sugar Yield

Parnidi - (  [email protected] )
1Research Group of Breeding on Sweetener, Tobacco, Fibre and Industrial Oil Crops, National Research and Innovation Agency Republic of Indonesia, Jl.
Raya Jakarta-Bogor KM 46, Cibinong, Jawa Barat 169
Weda Makarti Mahayu
Mala Murianingrum
Abdurrahman -
Sri Yulaikah
Djumali -
Bambang Heliyanto
Anik Herwati
Rully Dyah Purwati
Moch. Mahfud
Fatkhur Rochman
Marjani -
Taufiq Hidayat RS
2Research Group of Development of High Quality-Seed Production Technology for Horticulture and Estate Crops, National Research and Innovation
Agency Republic of Indonesia, Jl. Raya Jakarta-Bogor KM 46
Prima Diarini Riajaya
3Research Group of Precision Cultivation of Estate Crops, National Research and Innovation Agency Republic of Indonesia, Jl. Raya Jakarta-Bogor KM 46,
Cibinong, Jawa Barat 16911.
Fitriningdyah Tri Kadarwati
Budi Santoso
Supriyono -
4Research Group of Plant Pest-Diseases Detection System of Horticulture and Estate Crops, National Research and Innovation Agency Republic of
Indonesia, Jl. Raya Jakarta-Bogor KM 46, Cibinong, Jawa Ba
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Cece Suhara
4Research Group of Plant Pest-Diseases Detection System of Horticulture and Estate Crops, National Research and Innovation Agency Republic of
Indonesia, Jl. Raya Jakarta-Bogor KM 46, Cibinong, Jawa Ba
Research Article
Keywords:
Posted Date: November 9th, 2023
DOI: https://doi.org/10.21203/rs.3.rs-3570780/v1
License:   This work is licensed under a Creative Commons Attribution 4.0 International License. Read Full License
Additional Declarations: No competing interests reported.
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Abstract
Cane weight, sucrose content, and sugar yield represent carbohydrate accumulation available for cane throughout its life cycle. The carbohydrate content is
the result of photosynthesis; in other words, we can predict cane weight, sucrose content, and sugar yield from the photosynthesis process. Chlorophyll
fluorescence or the OJIP test is one of the benchmarks for expressing the amount of photosynthesis. This study aimed to obtain the dominant OJIP
variable as a predictor of cane weight, sucrose content, sugar yield, and measurement time. The study was conducted at the Asembagus Experimental
Station, Situbondo Regency, East Java, Indonesia, from December 2016 to October 2017. The study used 18 sugarcane clones in a Randomized Block
Design with 3 replications. OJIP variables were measured in the stalk elongation phase and the maturity phase. The results showed that OJIP variables
other than Fv/Fm, cane weight, sucrose content, and sugar yield were influenced by sugarcane clones. The right time for measuring OJIP variables was
during the maturity phase. The dominant OJIP variables as predictors of cane weight and sugar yield were TRo/RC, DIo/CS, ABS/RC, and PI (79.4 and
76.0%). The dominant predictors of yield were RC/CSo, RC/CSm, DIo/CS, PI, ABS/RC, and ETo/RC (92.9%).
1. Introduction
Cane weight and sucrose content make up the sugar yield-thus, the sugar yield can be predicted from the two components (Tyagi et al. 2012). Cane weight
is influenced by the carbohydrate available for stalk growth during the elongation and maturity phases (Jones et al.2011). During the cane elongation
phase, an increase in carbohydrates available for cane growth is followed by an increase in cane weight and vice versa (Khan et al. 2012).
The maturity phase of sugarcane means the storage of stored carbohydrates (sucrose) in the stalk—it can determine the sugar yield and cane weight
(Dashora 2012). The amount of stored carbohydrates is determined by the amount of carbohydrates available to the stalk during the maturity phase.
Carbohydrates available for stalks are used for stalk growth and stored carbohydrates. If sugarcane uses more carbohydrates for growth, it will hold less,
resulting in a lower yield. Therefore, during the maturity phase, sugarcane requires dry environmental conditions to produce a high yield (Cardozo et al.
2015).
From the explanation above, it can be concluded that the carbohydrates available for growth during the elongation and maturity phase can predict cane
weight, sucrose content, and sugar yield. Carbohydrates available for growth are the residue of photosynthesis after being used for respiration. Thus,
photosynthesis is the primary key to predicting cane weight, sucrose content, and sugar yield. Therefore, a decline in photosynthesis rates will reduce the
quantity of carbohydrates available for cane growth during the elongation phase and carbohydrates stored (sucrose) during the maturity phases (Zhao et
al. 2015).
Photosynthetic measurements are done in the CO2 fixation phase by calculating the amount of CO2 used in the photosynthesis process. Along with
technology development, photosynthetic measurements can now be done in the early phase of photosynthesis, called chlorophyll fluorescence. Chlorophyll
fluorescence or OJIP is a simple and non-invasive method for monitoring changes in photosynthetic processes by measuring the radiation emitted by
leaves (Yusuf et al. 2010). It is possible to calculate variables that can estimate energy absorption by antenna system pigments, exciton capture by reaction
center, and subsequent electron transport to the final electron acceptor (Stirbert et al. 2014). These measurements provide a constellation of structural and
functional variables that characterize the behavior of PS II (Khalid et al. 2015) and have been widely used to study PS II activity in various plants (Cuchiara
et al. 2013). For example, the on canola cultivars under salt stress (Stirbet 2011) use it on Alternanthera tenella colla under copper stress (Jedmowski et al.
2013) use it on Hordeum spontaneum and Sorghum bicolor under water stress, and use it on canola cultivars under light stress (Van Rensen and
Vredenberg 2011).
OJIP has many variables, yet there is no information on the dominant OJIP variables as predictors of cane weight, sucrose content, and sugar yield.
Likewise, there is no information on the measurement time, whether during the elongation or maturity phase. Therefore, this study was designed to reveal
the dominant OJIP variables as predictors of (a) cane weight, sucrose content, and sugar yield, and (b) measurement time. Our knowledge of these two
factors would help to determine the parents of a cross so that sugarcane crossbreeding can be done more efficiently and new high-yielding varieties can be
obtained in a shorter time.
2. Material and Methods

2.1 Study Site and Materials. The study was conducted at the Asembagus Experimental Station, Situbondo Regency, East Java, Indonesia, from December
2016 to October 2017. The soil type in the study site is Entisol. The physical and chemical properties of the soil are listed in Table 1. The rainfall during the
study is shown in Fig. 1.
The clones used were taken from a collection of sugarcane germplasm owned by the Indonesia Sweeteners and Fibers Crops Research Institute. Starting
June 2022, the Indonesian Sweetener and Fiber Crops Research Institute will be merged into National Research and Innovation Agency Republic of
Indonesia in accordance with Presidential Regulation No. 78 of 2021. We used two-eyed stem cutting from 18 sugarcane clones (17, 87, 90, 104, 212, 351,
354, 451, 452, PBG 2, 386 SOF 1118, PRG 881, MLG 19, 400 SOF 1172, 400 SOF 1132, PA 02.18, PS 881, and Cening). Other materials were inorganic and
organic fertilizers, pesticides, and other supporting materials. The tools included a refractometer, polarimeter, Chlorophyll Fluorometer type OS-30p+, and
other supporting tools.
2.2 Experiment Design and Culture Practices. The 18 sugarcane clones were arranged in a Randomized Block Design with 3 replications. OJIP variables
were measured in the stalk elongation phase and the maturity phase. Each clone in one replication was planted in 5 rows; each row was 5 meter-length. The
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center-to-center (CTC) distance was 130 cm. Before planting, we applied manure to each row with a dose of 10 t ha− 1. Each row was planted with 10
sugarcane stalks (stem cutting).
The maintenance of sugarcane included replanting, fertilizing, earthing-up, irrigating, and controlling pests and diseases. Replanting was done 2–3 weeks
after planting by planting available stem cuttings to replace the dead, damaged, or unhealthy stem cuttings planted; replanting aimed to ensure that the
plant population remained as planned. Fertilizers were applied twice: 1 and 3 months after planting. Fertilizers were applied to each row, approximately 10
cm from the stalk base. We used 750 kg Phonska and 625 kg Za per hectare. Phonska was applied on the first fertilization and Za on the second
fertilization. Earthing-up was done twice by piling up soil from the left and right of the row to the top of the row. It was done after the first and second
fertilization. Irrigation was applied 3 times from May to July.
The sugarcane was harvested 12 months after planting. First, all cane stalks with a minimum length of 150 cm and a minimum diameter of 2 cm on rows
2, 3, and 4 were cut from their base. Then, the stalks were cleaned from the dried leaves, and the top of the stalks was cut.
2.3 Cane Weight, Commercial Cane Sugar, and Sugar Yield Measurement. Cane weight was measured by weighting (CW) and counting the number of stalks
harvested on each plot (NS). Cane weight (CP) was measured with the following formula:
CW
CP =
NS
Sucrose content was observed by measuring fiber content, Brix, and Pol. We took a random sample of harvested cane stalks consisting of 6 stalks from
each block and each replication. The stalk sample was weighted (WS) and squeezed with a sample mill for its juice. The resulted juice was weighted (JW).
Brix of the sugarcane juice was measured using a hand refractometer, while Pol was measured using a polarimeter. The juice value (JV) is calculated with
the following formula:
J V = P ol − (0.4x (Brix − P ol))
Sucrose content (SC) is calculated with the following formula:
SC (%) = F P xN N
Sugar yield (SY) is measured with the following formula:
C P xSC
SY =
100
2.4 OJIP Variables Measurement. Observations of OJIP variables (chlorophyll fluorescence) were carried out in the elongation phase (5 months after
planting) and the maturity phase (10 months after planting). Chlorophyll fluorescence was measured with a Chlorophyll Fluorometer on fully-opened upper
leaves following the procedure for using the tool (Anonymous 2012). For each clone in one replication, we took 3 samples. Before measurement, the sample
leaves were conditioned in the dark for 30 minutes. The data recorded in the tool included the quantum yield of primary photochemistry PSII (Fv/Fm and
Fv/Fo), relative variable fluorescence at phase J of fluorescence transient curve (Vj), performance index (PI), the net rate of PS II closure ( Mo), the specific
flux (flux per active PSII reaction center) of absorption (ABS/RC), trapping (TRo/RC) and electron transport (ETo/RC), dissipation flux per excited cross-
section (DIo/CS), electron transport flux per excited cross-section (ETo/CS), the efficiency with which an exciton captured in the reaction center can move an
electron from QA− to the intersystem electron acceptor (ETo/TRo), and the density of the reaction center when all reaction centers are open (RC/CSo) and
when all PSII reaction centers are closed (RC/CSm).
2.5 Statistical Analysis. Data were analyzed for variance and continued with Duncan’s double distance test (DMRT) at a 5% significance level using MSTAT
software Version 4.00/EM. Multiple linear regression analysis (Stepwise analysis) between sugarcane weight, sucrose content, and sugar yield with OJIP
variables was done to determine how OJIP variables influenced the three agronomic variables. OJIP variables with an influence level > 5% were the
dominant variables influencing the three agronomic variables.
3. Results
3.1 Cane Weight, Commercial Cane Sugar, and Sugar Yield. Cane weight, commercial cane sugar, and sugar yield were affected by sugarcane clones (Table
2). Clones 17, 87, 104, 212, 354, 386 SOF1118, Cening, and PBG 2 produced the highest cane weight (1,444 to 1,647 kg stalk-1). The highest sucrose
content was found on PA 02.18, PRG 881, and PS 881 (11.66 to 11.86%). The highest sugar yield was obtained for 104, 386 SOF1118, and PS 881 (0.153-
0.164 kg stalk-1).
3.2 OJIP Variables. Sugarcane clones did not affect Fv/Fm but affected other OJIP variables when measured during the elongation and maturity phase
(Table 3 and 4). The highest Fv/Fo measured during the elongation phase came from clones 17, 90, 351, 400 SOF1172, 400 SOF1132, PA 02.18, PBG 2,
PRG 881, and PS 881. The highest Fv/Fo measured during the maturity phase came from clones 351, 386 SOF1118, PA 02.18, and PS 881.
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The highest Vj measured during the elongation phase came from clones Cening, MLG 19, PBG 2, and PRG 881. The highest Vj measured during the maturity
phase came from clones 400 SOF1132, 400 SOF1172, Cening, and MLG 19.
The highest PI measured during the elongation phase came from clones 90 and PS, while during the maturity phase came from clones 351, 452, and PBG
2.
The highest Mo measured during the maturity phase came from clone Cening, while during the maturity phase came from clone MLG 19.
The highest ABS/RC measured during the elongation phase came from clones 17, 104, 212, 351, 354, 451, 400 SOF1172, and Cening, while during the
maturity phase came from clones 17, 354, and MLG 19.
The highest TRo/RC measured during the elongation phase came from clones 212 and Cening, while during the maturity phase came from clones 17, 104,
354, Cening, MLG 19, PA 02.18, and PS 881.
The highest ETo/RC measured during the elongation phase came from clones 104, 351, and 451, while during the maturity phase came from clone 354.
The highest DIo/CS measured during the elongation phase came from clones 104, 212, 354, 451, 400 SOF1172, Cening, and MLG 19, while during the
maturity phase came from clone 17.
The highest ETo/TRo measured during the elongation phase came from clones 90, 104, 351, 354, 451, 386 SOF1118, and PS 881, while during the maturity
phase came from clones 90, 351, 354, 452, 386 SOF1118, PA 02.18, and PS 881. The highest ETo/CS measured during the elongation phase came from
clones 90, 104, 351, 451, and PS 881, while during the maturity phase came from clone 452.
The highest RC/CSo measured during the elongation phase came from clones 90 and 452, while during the maturity phase came from clones 87 and
351. The highest RC/CSm measured during the elongation phase came from clone 90, while during the maturity phase came from clone 351.
3.3 The Relationship of OJIP variables with Cane Weight, Sucrose Content, and Sugar Yield. The stepwise analysis between sugarcane weight and OJIP
variables resulted in a correlation coefficient of 0.667 in the elongation phase and 0.863 in the maturity phase (Table 5). These results mean that the 12
OJIP variables affected cane weight with a total effect of 66.70% in the elongation phase and 86.33% in the maturity phase. Thus, it can be concluded that
the appropriate time for measuring the OJIP variable to predict the cane weight was during the maturity phase.
Vj, Mo, TRo/RC, DIo/CS, ETo/TRo, ETo/CS, and RC/CSm positively affected cane weight during the elongation phase, while other variables negatively
affected cane weight. Fv/Fo, ABS/RC, and RC/Cso positively affected cane weight during the maturity phase, while other variables negatively affected cane
weight. During the elongation phase, RC/CSo, ETo/CS, Mo, DIo/CS, and ETo/RC became the dominant OJIP variables affecting cane weight with a total
effect of more than 50%. During the maturity phase, TRo/RC, DIo/CS, ABS/RC, and PI became the dominant OJIP variables affecting cane weight with a
total effect of more than 70%.
The stepwise analysis of sucrose content resulted in a correlation coefficient of 0.800 during the elongation phase and 0.979 during the maturity phase
(Table 6). During the elongation phase, Fv/Fo, Vj, PI, Mo, ABS/RC, and ETo/CS positively affected sucrose content, while other variables negatively affected
sucrose content. Variables with the most dominant effect, with a total effect of 80%, were, from the highest effect to the lowest, RC/CSm, RC/CSo, Fv/Fo,
TRo/RC, PI, DIo/CS, and ABS/RC, respectively. During the maturity phase, Fv/Fo, ABS/RC, and RC/CSm positively affected sucrose content, while other
variables negatively affected sucrose content. The OJIP variables with the most dominant effect, with a total effect of more than 75%, were, from the
highest effect to the lowest, RC/CSo, RC/CSm, DIo/CS, dan PI, respectively. Thus, the best time to measure OJIP variables to predict sucrose content was
during the maturity phase.
The analysis of the relationship of sugar yield with OJIP variables resulted in a correlation coefficient of 0.729 during the elongation phase and 0.847
during the maturity phase (Table 7). Thus, the 12 OJIP variables affected sugar yield with a total effect of 72.90% during the elongation phase and 84.71%
during the maturity phase. TRo/RC, DIo/CS, ETo/TRo, ETo/RC, and RC/CSm negatively affected sugar yield during the elongation phase, while Fv/Fo, Mo,
ABS/RC, and RC/CSm positively affected sugar yield during the maturity phase. The OJIP variables measured during the maturity phase contributed higher
to sugar yield than during the elongation phase. Thus, the best time to measure OJIP variables to predict sugar yield was during the maturity phase.
The OJIP variables with the most dominant effect on sugar yield during the elongation phase, with a total effect of more than 60%, were RC/CSm, PI,
ETo/CS, ETo/RC, and PI. The OJIP variables with the most dominant effect on sugar yield during the maturity phase, with a total effect of more than 70%,
were, from the highest effect to the lowest, TRo/RC, DIo/CS, ABS/RC, and PI.
4. Discussion
4.1 Cane Weight, Sucrose Content, and Sugar Yield. Interaction between environmental conditions and clones affects cane weight, sucrose content, and
sugar yield (Kumar et al. 2012). If the environmental conditions are homogeneous, the three agronomic variables are influenced by clones. We used
homogenous environmental conditions in this present study, so differences in results were due to the clones used. The differences in cane weight, sucrose
content, and sugar yield are due to differences in the clones used (Chohan et al. 2014; Ahmed 2017 and Schultz et al. 2017).
4.2 OJIP Variables. The maximum quantum yield of primary photochemistry (Fv) is standardized with the values of Fm and Fo, so we obtain Fv/Fm = (Fm-
Fo)/Fm and Fv/Fo = (Fm-Fo)/Fo. The Fm value was relatively high (485.11 to 671.17) in the elongation phase and 608.44 to 760.50 in the maturity phase,
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while the Fo value was relatively low (144.67 to 211.78) in the elongation phase and 148.56–174.50 in the maturity phase. The significant differences in
the value of Fm and Fo caused the Fv/FM values to be less diverse, while the Fv/Fo values to be diverse. This condition caused Fv/Fm to be unaffected by
the clones, while the Fv/Fo was affected by the clones. Studies on sugarcane under aluminum stress (Ecco et al. 2013) and on Alternanthera tenella colla
under copper stress and sweet potato under copper stress also slow low diversity of Fv/Fm values and diverse Fv/Fo values (Fghire et al. 2015). However,
the study on Chenopodium quinoa shows different Fv/Fm values due to water stress (Figuiredo et al. 2015).
The morphology of sugarcane leaves, including the number of stomata, the number of epidermal cells, the polar diameter of the stomata (stomata length),
the equatorial diameter of the stomata (stomata width), the thickness of the epidermis on the lower and upper surfaces, the thickness of the mesophyll, the
thickness of the upper cuticle, the polar diameter of the bulliform cells, the number of bulliform cells, the diameter of the bundle sheath cells, the thickness
of the phloem, the number of metaxylem vessels, the diameter of the metaxylem vessels, and the distance between the vascular bundles, are affected by
the clone (Khan et al. 2017). In addition, leaf color, wax layer thickness, leaf hair density, and sugarcane leaf thickness are influenced by the clones used
(Chidambaram et al. 2017 and Martinazzo et al. 2012). Differences in leaf morphology cause differences in the amount of light received, reflected, and
absorbed by the leaves (Desotgiu et al. 2012). Such conditions cause the Fv/Fo, Vj, PI, and Mo values produced by each sugarcane clone to differ, both in
the stem elongation and maturity phases. Two sugarcane clones respond differently to different water availability and aluminum levels in producing Fv/Fo,
Vj, PI, and Mo values (Chohan et al. 2014).Different leaf thickness and chlorophyll content in leaves affect the number of reaction centers (Pedrozo et al.
2015 and Tadesse et al. 2014). The thicker the leaf and the higher the chlorophyll content, the more reaction centers the leaf has. Leaf thickness and
chlorophyll content in leaves are one of the characteristics of sugarcane clones (Tena et al. 2016; Avivi et al. 2019 and Beneragama et al. 2014). This
condition causes RC/CSo and RC/CSm to be affected by the sugarcane clones.
The absorption energy (ABS) will be absorbed (TR)—the energy will be partly used for electron transport (ET) and partly lost as heat energy (DIo) (Farias et
al. 2016). Different ABS/RC values are caused by differences in antennas in the photosystem complex (Li et al. 2012). The difference in ABS/RC causes
differences in the values of DIo/RC, TRo/RC, ETo/RC, and ETo/TRo obtained (Jedmowski et al. 2015 and Kalaji et al. 2011). The difference in antennas in
the photosystem complex is caused by differences in the cultivars used (Kalaji et al. 2011). This causes the sugarcane clones to influence the ABS/RC,
DIo/CS, TRo/RC, ETo/RC, ETo/TRo, and ETo/CS values. The different canola cultivars result in different ABS/RC, DIo/CS, TRo/RC, and ETo/RC values
(Stirbert et al. 2014).
4.3 The Relationship of OJIP Variables with Cane Weight, Commercial Cane Sugar, and Sugar Yield. Harvested cane weight is the accumulation of (1)
carbohydrates available for cane growth during the elongation and maturity phase and (2) carbohydrates stored (sucrose) during the maturity phases.
Sugarcane keeps carbohydrates in the stem tissue as sucrose. Carbohydrates available for growth are the residue of photosynthesis after being used for
respiration. Fluorescence chlorophyll (OJIP variable) analysis provides quick insight into the ability of plants to photosynthesize to tolerate environmental
pressure (Kalaji et al. 2011). This condition caused the OJIP variables observed in the elongation phase to contribute significantly (82.34%) in influencing
sugarcane weight, yet it contributed only 73.20% and 71.72% in influencing sucrose content and sugar yield. The OJIP variables observed in the maturity
phase contributed 86.33%, 97.88%, and 84.71% in influencing sugarcane weight, sucrose content, and sugar yield, respectively. Thus, observing OJIP
variables in the maturity phase is more appropriate for predicting sugarcane productivity, sucrose content, and sugar yield.
The light energy the leaves receive is absorbed by the chlorophyll, and some of the energy is reflected. The absorbed energy (ABS) partially undergoes
adsorption (TR), and the rest turns into heat and fluorescent energy (dissipate = DI) (Beneragama et al. 2014). The absorbed energy is then used for electron
transport (ET). In general, an increase in the light energy absorbed in chlorophyll causes an increase in the photosynthesis rate so that carbohydrates are
available for growth and storage. This condition causes ABS/RC to positively affect cane weight, sucrose content, and sugar yield. Likewise, the higher the
dissipated energy (DIo/RC), the lower the energy used for electron transport (ETo/RC), so DIo/CS negatively affects cane weight, sucrose content, and sugar
yield.
In stepwise analysis, Xn negatively affects Y, which can have two meanings. First, it means the individual influence of Xn on Y is negative. Second, it means
the individual influence of Xn on Y is positive, but the positive value is below the positive value of the combined effect of the X value. In the case of TRo/RC,
which negatively affects cane weight and sugar yield, the second meaning applies. The TRo/RC value is the reduction of the ABS/RC value with the DIo/CS
value. The ABS/RC has a positive effect, and DIo/CS has a negative effect, so TRo/RC individually has a positive effect on the two agronomic variables
(Beneragama et al. 2014).
The Performance Index (PI) describes the overall expression of the plant’s internal strength to deal with environmental conditions. PI depends on the three
functional stages of photosynthetic activities by the RC PSII complex (light energy absorption, excitation energy absorption, and the conversion of absorbed
energy to electron transport in PSII) (Strasser et al. 2004). In this study, the excitation energy absorption (TRo/RC) and the conversion of the adsorbed
energy to electron transport (ETo/RC) negatively affected cane weight, sucrose content, and sugar yield—thus, PI negatively affected the three agronomic
variables.
5. Conclusion
The OJIP variables, other than Fv/Fm, observed in the elongation and maturity phase were influenced by sugarcane clones. Our findings confirmed that the
maturity phase was the best time for measuring OJIP variables to predict cane weight, sucrose content, and sugar yield. The dominant OJIP variables as
predictors of cane weight and sugar yield were TRo/RC, DIo/CS, ABS/RC, and PI, with an accuracy of 79.4% and 76.0%, respectively. The dominant
predictors of sugar yield were RC/CSo, RC/CSm, DIo/CS, PI, ABS/RC, and ETo/RC, with an accuracy of 92.9%.
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Declarations
Data Availability
Data are available upon request from the corresponding author by interviews with expert staff at the office of National Research and Innovation Agency
Republic of Indonesia, Jl. Raya Jakarta-Bogor KM 46, Cibinong, Jawa Barat 16911.
Conflicts of Interest
The authors declare that they do not have any commercial or associative interest that represents a confict of interest in connection with the work
submitted.
Author contributions
Parnidi (PR), Weda Makarti Mahayu (W.M), Mala Murianingrum (M.M), Abdurrahman (AB), Sri Yulaikah (S.Y), Djumali (DJ), Bambang Heliyanto (B.H), Anik
Herwati (A.H), Rully Dyah Purwati (R.D), Moch. Mahfud (Mo.M), Fatkhur Rochman (F.R), Marjani (MJ), Taufiq Hidayat RS (T.H), Prima Diarini Riajaya (P.D),
Fitriningdyah Tri Kadarwati (F.K), Budi Santoso (B.S), Supriyono (SP), and Cece Suhara (C.S) proposed the study and concept of work. Parnidi (PR), Taufiq
Hidayat RS (T.H), Weda Makarti Mahayu (W.M), Mala Murianingrum (M.M), and Abdurrahman (AB). collected data and cleaned data. Djumali (DJ), Marjani
(MJ) and Fatkhur Rochman (F.R). interpreted the data. Djumali (DJ), Parnidi (PR) and Taufiq Hidayat RS (T.H). analyzed the results. Supriyono (SP), Cece
Suhara (C.S), and Moch. Mahfud (Mo.M). wrote the paper. Marjani (MJ) and Fatkhur Rochman (F.R), Anik Herwati (A.H), Rully Dyah Purwati (R.D), Sri
Yulaikah (S.Y). reviewed and edited the paper. Djumali (DJ), Mala Murianingrum (M.M), Weda Makarti Mahayu (W.M) and Rully Dyah Purwati (R.D). made
critical revisions of the article. Supriyono (SP), Cece Suhara (C.S), and Moch. Mahfud (Mo.M), Bambang Heliyanto (B.H), Prima Diarini Riajaya (P.D), and
Fitriningdyah Tri Kadarwati (F.K). supervised the process. All authors have read and agreed to the published version of the manuscript. All the authors
contributed equally to this research and drafting this manuscript.
Acknowledgments
We thank the Head of the Indonesia Sweetener and Fiber Crops Research Institute for funding this research through the 2017 DIPA. Starting June 2022, the
Indonesian Sweetener and Fiber Crops Research Institute will be merged into National Research and Innovation Agency Republic of Indonesia in
accordance with Presidential Regulation No. 78 of 2021. We also thank the Head of the Asembagus Experimental Station and all parties who made this
study possible.
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the-fluorescence-transient.pdf
Tables
Table 1. Physical and chemical characteristics of soil in the study site (Asembagus Experimental Station)
Page 8/13
Soil characteristics Value Categories
pH 1:1 H2O 7.0 Neutral
pH KCl 1 N 6.8
C-Organic (%) 0.56 Very low
N-total (%) 0.06 Very low
C/N 9,3 Low
P Olsen (mg kg-1) 51.38 Very high
K (NH4OAC 1N pH:7) (me/100 g) 1.42 Very high
Na (NH4OAC 1N pH:7) (me/100 g) 1.09 Very high
Ca (NH4OAC 1N pH:7) (me/100 g) 11.58 High
Mg (NH4OAC 1N pH:7) (me/100 g) 1.81 Medium
CEC (me/100 g) 15.87 Low
Total bases 15.90
Base Saturation (%) 100 Very high
Sand (%) 81
Silk (%) 19 Loamy sand
Clay (%) 0
Table 2. Cane Weight, Commercial Cane Sugar, and Sugar Yield of Sugarcane Clones
Clones Cane weight Commercial Cane Sugar (%) Sugar yield
(kg stalk-1) (kg stalk-1)
17 1.617 a 6.07 g 0.098 gh
87 1.560 a 6.33 fg 0.102 f-h
90 1.338 b-d 6.25 f-g 0.085 h
104 1.629 a 9.43 bc 0.153 ab
212 1.617 a 8.31 c-e 0.134 bc
351 1.198 d 10.17 b 0.122 c-f
354 1.600 a 7.74 d-f 0.126 cd
451 1.213 d 8.00 c-e 0.097 gh
452 1.000 e 9.39 bc 0.093 h
386 SOF1118 1.647 a 9.91 b 0.164 a
400 SOF1132 1.247 cd 9.85 b 0.125 c-e
400 SOF1172 1.355 b-d 7.71 d-f 0.104 e-h
Cening 1.481 ab 6.94 e-g 0.105 d-h
MLG 19 1.276 cd 9.16 b-d 0.116 c-g
PA 02.18 0.978 e 11.86 a 0.116 c-g
PBG 2 1.444 a-c 9.46 bc 0.136 bc
PRG 881 0.899 e 11.69 a 0.104 e-h
PS 881 1.358 b-d 11.66 a 0.159 a
Note:
Page 9/13
1. Values in the same column followed by the same letters were not significantly different at a 5% significance level based on the Duncan Multiple Range
Test
2. NS = not significantly differed
Table 3. The maximum photochemical quantum yield of PSII (Fv/Fm and Fv/Fo), relative variable fluorescence at phase J of fluorescence transient curve
(Vj), performance index (PI) and net rate of PS II closure (Mo), the specific flux (flux per active PSII reaction centre) of absorption (ABS/RC), trapping
(TRo/RC) and electron transport (ETo/RC), the specific flux of dissipation per excited cross section (DIo/CS), the efficiency with which an exciton captured
in the reaction centre can move an electron from QA- to the intersystem electron acceptor (ETo/TRo) and electron transport per excited cross section
(ETo/CS), densities of reaction center when all reaction centers opened (RC/CSo) and when all reaction centers PSII closed (RC/CSm) of of sugarcane
clones on stalk elongation phase.
Clones Fv/Fm Fv/Fo Vj PI Mo ABS/RC TRo/RC ETo/RC DIo/CS ETo/TRo ETo/CS RC/CSo RC/CSm
17 0.759ns 3.19 a- 0.349 5.35 0.303 1.143 a- 0.196 0.564 0.276 0.651 b- 81.23 143.8 c- 605.1 d-
c d-f d-g b-f c bc c-e b-f d cd e f
87 0.742 2.94 b- 0.338 5.60 0.265 1.060 0.175 c- 0.521 0.274 0.662 b- 83.58 160.6 b- 631.4 c-
d d-f d-f d-g cd f d-f b-f d b-d d e
90 0.770 3.36 ab 0.271 11.03 a 0.196 0.920 0.138 f 0.513 0.211 0.729 ab 98.03 a 192.2 a 837.8 a
hi h de d-f g
104 0.743 2.96 b- 0.268 0.248 1.269 0.181 0.699 a 0.322 0.732 ab 94.45 138.3 ef 542.8 e-
d hi 6.59 b- f-h ab cd ab ab g
d
212 0.728 2.74 cd 0.368 3.99 0.349 1.290 a 0.218 0.590 0.352 0.632 c- 80.17 138.3 ef 517.3 fg
c-e g-i bc ab b-d a e cd
351 0.760 3.18 a- 0.275 0.244 1.187 a- 0.177 c- 0.658 0.286 0.725 ab 92.23 144.6 c- 606.8 d-
c g-i 7.25 bc f-h c e ab b-e a-c e f
354 0.730 2.75 cd 0.302 0.252 1.151 a- 0.175 c- 0.584 0.315 0.698 a- 85.61 150.4 c- 568.8 d-
f-i 5.73 d- f-h c f b-e ab c bc e f
f
451 0.744 2.98 b- 0.277 7.61 b 0.246 1.191 a- 0.176 c- 0.630 0.316 0.723 ab 89.79 151.4 c- 618.5 c-
d g-i f-h c e a-c ab a-c e e
452 0.735 2.85 cd 0.381 5.98 0.230 0.829 e 0.142 0.379 g 0.220 0.619 c-f 73.36 193.9 a 749.2 b
b-d c-f gh ef fg de
386 0.746 2.98 b- 0.319 0.249 1.049 0.170 c- 0.533 0.267 0.681 a- 85.52 160.5 b- 637.2 cd
SOF d e-h 6.18 c- f-h cd f d-f b-g c bc d
1118 e
400 0.749 3.03 a- 0.333 7.19 0.258 1.009 c- 0.168 c- 0.496 0.256 0.667 b- 82.37 172.9 b 702.9 bc
SOF d d-g bc e-h e f ef c-g d cd
1132
400 0.750 3.06 a- 0.331 5.53 0.298 1.199 a- 0.198 0.598 0.303 0.669 b- 84.90 141.9 de 580.4 d-
SOF c d-g d-f c-g c bc b-d a-d d b-d f
1172
CENING 0.736 2.85 cd 0.432 3.20 i 0.415 1.305 a 0.232 a 0.541 0.348 0.568 ef 66.41 122.2 f 475.2 g
ab a c-f a ef
MLG 19 0.718 2.59 d 0.460 0.366 1.073 b- 0.186 0.400 g 0.307 0.540 f 61.53 f 155.3 b- 561.6 d-
a 3.44 hi ab d bc a-c e f
PA 0.749 3.03 a- 0.343 0.270 1.072 b- 0.175 c- 0.525 0.277 0.657 b- 84.96 161.3 b- 636.5 cd
02.18 d d-f 5.85 c-f d-g d f d-f b-f d b-d d
PBG 2 0.760 3.18 a- 0.412 4.53 f- 0.321 1.030 0.188 0.461 0.248 0.588 d-f 67.78 148.3 c- 620.6 c-
c a-c i b-e cd bc fg d-g ef e e
PRG 0.758 3.17 a- 0.411 4.78 0.326 1.039 0.185 0.462 0.251 0.589 d-f 66.54 143.1 de 599.3 d-
881 c a-c e-h b-d cd bc fg c-g ef f
PS 881 0.777 3.48 a 0.247 10.57 a 0.193 1.026 0.146 0.605 0.228 0.753 a 94.70 164.1 bc 732.0 b
i h cd d-f b-d e-g ab
Values in the same column followed by same letters were not significantly different at 5% level base on the Duncan Multiple Range Test. NS = non
significant.
Table 4. The maximum photochemical quantum yield of PSII (Fv/Fm and Fv/Fo), relative variable fluorescence at phase J of fluorescence transient curve
(Vj), performance index (PI) and net rate of PS II closure (Mo), the specific flux (flux per active PSII reaction centre) of absorption (ABS/RC), trapping
(TRo/RC) and electron transport (ETo/RC), the specific flux of dissipation per excited cross section (DIo/CS), the efficiency with which an exciton captured
in the reaction centre can move an electron from QA- to the intersystem electron acceptor (ETo/TRo) and electron transport per excited cross section
Page 10/13
(ETo/CS), densities of reaction center when all reaction centers opened (RC/CSo) and when all reaction centers PSII closed (RC/CSm) of sugarcane clones
on maturity phases
Clones Fv/Fm Fv/Fo Vj PI Mo ABS/RC TRo/RC ETo/RC DIo/CS ETo/TRo ETo/CS RC/CSo RC/CSm
17 0.617 1.88 0.483 2.52 gh 0.488 1.713 a 0.229 a 0.483 0.742 0.517 cd 67.45 137.7 427.8 hi
NS gh bc b b-d a e-g gh
87 0.633 1.85 0.398 5.70 b- 0.231 0.939 0.132 f- 0.316 0.392 0.602 b- 74.84 302.4 a 893.6 b
gh de d e-g ef h fg de c c-e
90 0.613 1.64 h 0.355 2.38 gh 0.274 1.343 b- 0.174 0.551 b 0.518 0.645 ab 80.81 166.0 e- 435.6 hi
e d-f d b-e c b-d g
104 0.685 2.20 e- 0.486 3.40 fg 0.414 1.157 c- 0.192 0.376 0.367 0.514 cd 60.35 189.0 d- 605.9 e-
g bc bc e a-d ef d-f gh f g
212 0.670 2.28 c- 0.400 3.40 fg 0.273 1.136 c- 0.166 c- 0.459 0.404 0.600 bc 76.11 196.0 de 645.2 ef
g de d-f e f c-e de b-e
351 0.728 2.70 0.358 7.49 a 0.191 0.680 0.117 0.307 0.181 i 0.642 ab 80.86 326.7 a 1184.4
a-c e fg fg gh fg b-d a
354 0.648 1.98 f- 0.343 4.15 ef 0.398 1.778 a 0.210 0.732 a 0.648 0.657 ab 85.80 112.0 h 344.7 i
h ef bc ab b b
451 0.652 1.91 f- 0.421 3.26 fg 0.292 0.998 0.156 0.350 0.356 0.579 bc 67.08 225.5 b- 672.3 d-
h c-e de ef d-f fg d-g e-g d f
452 0.676 2.09 f- 0.278 6.50 ab 0.176 1.016 0.130 f- 0.512 0.328 0.722 a 101.38 250.3 bc 769.4 cd
h f g de h bc e-g a
386 0.710 2.66 0.388 5.84 bc 0.274 0.957 0.149 e- 0.400 0.283 0.612 a- 71.91 177.5 e- 650.5 ef
SOF a-d de d-f ef g d-f gh c d-f g
1118
400 0.699 2.36 c- 0.543 2.39 gh 0.411 1.019 0.174 0.312 0.296 0.457 de 45.96 164.3 e- 527.6
SOF f ab bc de b-e fg f-h i g gh
1132
400 0.646 1.94 f- 0.609 1.62 h 0.464 1.152 c- 0.151 e- 0.258 g 0.429 0.391 e 46.38 172.8 e- 530.0
SOF h a b e g d i g gh
1172
CENING 0.642 1.96 f- 0.541 1.46 h 0.478 1.399 0.211 0.392 0.528 0.459 de 54.29 137.8 412.8 i
h ab b bc ab ef c hi gh
MLG 19 0.672 2.23 0.608 1.53 h 0.643 1.580 0.212 a 0.386 0.534 0.392 e 44.54 111.3 h 367.5 i
d-g a a ab ef c i
PA 0.754 3.07 a 0.379 5.23 cd 0.336 1.165 c- 0.198 0.541 0.289 0.621 a- 77.50 157.6 e- 647.2 ef
02.18 e cd e a-c bc f-h c b-e g
PBG 2 0.656 2.36 c- 0.460 6.94 a 0.194 0.616 g 0.110 h 0.262 g 0.160 i 0.540 b- 62.40 218.3 cd 699.6 de
f cd fg d f-h
PRG 0.718 2.63 0.458 4.71 de 0.284 0.833 e- 0.151 e- 0.329 0.219 0.542 b- 60.20 257.9 b 854.7 bc
881 b-e cd d-f g g fg hi d gh
PS 881 0.739 2.85 0.345 4.72 de 0.293 1.154 c- 0.192 0.558 b 0.303 0.655 ab 84.69 152.2 fg 586.6 fg
ab ef de e a-d fg bc
Values in the same column followed by same letters were not significantly different at 5% level base on the Duncan Multiple Range Test. NS = non
significant.
Table 5. Regression Coefficients and Contribution Values on the Relationship of OJIP Variables with Cane Weight on the Elongation and Maturity Phases.
Page 11/13
OJIP variables Elongation Phase Maturity Phase
Regression Coefficient Contribution Values (%) Regression Coefficient Contribution Values (%)
Fv/Fo -0.494 1.76 0.421 1.51
Vj 1.652 0.00 -3.486 0.61
PI -0.583 3.53 -0.679 5.17
Mo 3.307 5.64 -0.782 0.30
ABS/RC -7.722 4.58 19.314 19.15
TRo/RC 0.940 0.35 -11.832 35.25
DIo/CS 2.840 5.29 -6.868 19.79
ETo/RC -4.625 5.29 -1.300 0.61
ETo/TRo 5.636 0.35 -3.595 0.00
ETo/CS 1.915 16.57 -0.376 0.61
RC/CSo -1.722 22.92 0.315 0.91
RC/CSm 7.195 2.09 -0.724 2.43
Intercept -5.740 8.835
Correlation coefficient 0.677 0.863

Total contribution 67.70 86.33
Table 6. Regression Coefficients and Contribution Values on the Relationship of OJIP Variables with Sucrose Content on the Elongation and Maturity
Phases
OJIP variables Elongation phase Maturity phase
Regression Coefficient Contribution Values Regression Coefficient Contribution Values

(%) (%)
Fv/Fo 1.452 10.08 0.323 3.31
Vj 0.286 0.00 -1.198 0.00
PI 1.069 7.73 -0.456 9.94
Mo 0.222 0.00 1.711 0.00
ABS/RC 11.435 6.72 6.158 8.28
TRo/RC -8.012 9.75 -1.388 1.65
DIo/CS -3.942 7.06 -2.571 11.60
ETo/RC -0.432 0.00 -2.459 8.28
ETo/TRo -4.974 0.00 0.408 0.00
ETo/CS 0.101 0.00 0.056 0.00
RC/CSo 1.912 18.82 -0.956 34.92
RC/CSm -2.644 19.83 0.976 19.93
Intercept 4.411 1.430

Table 7. Regression Coefficients and Contribution Values on the Relationship of OJIP Variables with Sugar Yield on the Elongation and Maturity Phases
Page 12/13
OJIP variables Elongation Phase Maturity Phase
Regression Coefficient Contribution Values Regression Coefficient Contribution Values
(%) (%)
Fv/Fo 0.436 1.94 0.362 0.58
Vj 1.608 0.00 -5.300 0.29
PI 0.793 10.97 -1.233 8.83
Mo 1.830 2.58 3.479 0.45
ABS/RC 5.065 3.23 27.501 20.29
TRo/RC -5.936 12.90 -13.002 23.51
DIo/CS -1.624 2.58 -10.417 23.51
ETo/RC -2.650 2.58 -5.149 3.78
ETo/TRo -1.138 0.00 -2.801 0.14
ETo/CS 1.471 16.13 -0.126 0.14
RC/CSo 0.116 0.00 -0.817 2.90
RC/CSm -1.705 20.00 0.342 0.29
Intercept 0.085 9.845

Figures
Figure 1
Rainfall during the study
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Derivate OJIP Variables in Sugarcane to Predict Cane Weight, Sucrose

Content, and Sugar Yield

Posted Date: November 9th, 2023

Additional Declarations: No competing interests reported.

2. Material and Methods

J V = P ol − (0.4x (Brix − P ol))

Sucrose content (SC) is calculated with the following formula:

Sugar yield (SY) is measured with the following formula:

pH 1:1 H2O 7.0 Neutral

C-Organic (%) 0.56 Very low

N-total (%) 0.06 Very low

C/N 9,3 Low

P Olsen (mg kg-1) 51.38 Very high

K (NH4OAC 1N pH:7) (me/100 g) 1.42 Very high

Na (NH4OAC 1N pH:7) (me/100 g) 1.09 Very high

Ca (NH4OAC 1N pH:7) (me/100 g) 11.58 High

Mg (NH4OAC 1N pH:7) (me/100 g) 1.81 Medium

CEC (me/100 g) 15.87 Low

Total bases 15.90

Base Saturation (%) 100 Very high

Silk (%) 19 Loamy sand

Clones Cane weight Commercial Cane Sugar (%) Sugar yield

(kg stalk-1) (kg stalk-1)

17 1.617 a 6.07 g 0.098 gh

87 1.560 a 6.33 fg 0.102 f-h

90 1.338 b-d 6.25 f-g 0.085 h

104 1.629 a 9.43 bc 0.153 ab

212 1.617 a 8.31 c-e 0.134 bc

351 1.198 d 10.17 b 0.122 c-f

354 1.600 a 7.74 d-f 0.126 cd

451 1.213 d 8.00 c-e 0.097 gh

452 1.000 e 9.39 bc 0.093 h

386 SOF1118 1.647 a 9.91 b 0.164 a

400 SOF1132 1.247 cd 9.85 b 0.125 c-e

400 SOF1172 1.355 b-d 7.71 d-f 0.104 e-h

Cening 1.481 ab 6.94 e-g 0.105 d-h

MLG 19 1.276 cd 9.16 b-d 0.116 c-g

PA 02.18 0.978 e 11.86 a 0.116 c-g

PBG 2 1.444 a-c 9.46 bc 0.136 bc

PRG 881 0.899 e 11.69 a 0.104 e-h

PS 881 1.358 b-d 11.66 a 0.159 a

Fv/Fo -0.494 1.76 0.421 1.51

Vj 1.652 0.00 -3.486 0.61

PI -0.583 3.53 -0.679 5.17

Mo 3.307 5.64 -0.782 0.30

ABS/RC -7.722 4.58 19.314 19.15

TRo/RC 0.940 0.35 -11.832 35.25

DIo/CS 2.840 5.29 -6.868 19.79

ETo/RC -4.625 5.29 -1.300 0.61

ETo/TRo 5.636 0.35 -3.595 0.00

ETo/CS 1.915 16.57 -0.376 0.61

RC/CSo -1.722 22.92 0.315 0.91

RC/CSm 7.195 2.09 -0.724 2.43

Intercept -5.740 8.835

Correlation coefficient 0.677 0.863

OJIP variables Elongation phase Maturity phase

Regression Coefficient Contribution Values Regression Coefficient Contribution Values

Fv/Fo 1.452 10.08 0.323 3.31

Vj 0.286 0.00 -1.198 0.00

PI 1.069 7.73 -0.456 9.94