Agrarian Academic Journal
doi: 10.32406/v7n3/2024/27-42/agrariacad
Effects of temperature fluctuations on the germination and vigor of durum wheat seeds in Algeria. Efeitos das flutuações de temperatura na germinação e no vigor de sementes de trigo duro na Argélia.
Manel Lina Djendi
1, Amel Soussa2*, Nabila Bouhaddouda3, Amina Dridi2, Ibtissem Zeriri2, Hiba Daas2
1- Laboratory of Microbiology and Molecular Biology, Department of Biochemistry, Faculty of Sciences, Badji-Mokhtar – Annaba University, Box 12, Annaba 23000, Algeria. E-mail: linadjendi23@gmail.com
2– Environmental Research Center, Alzon Castle, Boughazi Said Street, PB 2024, Annaba 23000, Algeria. E-mail: soussa-amel@hotmail.fr, bilamina@hotmail.com, zeririibtissem@gmail.com, hiba_daas@outlook.fr
3- Faculty of Science, Biology Department, Skikda, Algeria. E-mail: bouhaddouda.n@hotmail.fr
*- Correspondence author. E-mail: soussa-amel@hotmail.fr
Abstract
Understanding the influence of temperature variations on durum wheat seed germination is of practical importance to farmers, researchers and agricultural decision-makers. The ability of seeds to germinate optimally is crucial to ensuring satisfactory crop yields. The main objective of this work is to analyze the impact of temperature variations on the viability and vigor of durum wheat seeds in Algeria. To achieve this goal, we adopted a comprehensive approach that primarily focused on studying the morphological, physiological, biochemical, and technological parameters (such as germination, seedling growth kinetics, soluble protein, proline, and soluble sugar content) of the seeds. Indeed, over a span of two years, four wheat varieties (Bousselam, Setifis, Megress, Saoura) were subjected to two distinct temperature and humidity conditions for the duration of the study. The first condition involved ambient temperatures fluctuating between 11°C and 37°C, with relative humidity ranging from 30% to 85%. However, the second condition involved storing the seeds in a cold room at a temperature of -04°C and a humidity of 12%. The results show significant variations in the germination rate of all varieties under both temperature conditions F=40, 54 P=0,000, with variety Broving to be the most resistant to thermal changes. According to the results obtained, temperature affects the growth of the roots of the S variety, while the Bousselam (B), Megress(M), Saoura (SW)re not affected; in addition, storage under both temperature conditions has an impact on the average number of roots and the length of coleoptiles. Protein analyses revealed significant effects of storage under ambient conditions (F=4,70 P=0,000), while in the cold room, these effects were attenuated. In addition, storage showed variations in soluble sugar and proline content, with different responses under the two temperature conditions. These results underline the importance of understanding the responses of durum wheat varieties to variable thermal conditions, offering useful prospects for improving agricultural practices and preserving seed quality in changing environments.
Keywords: Temperature. Germination. Durum wheat. Seeds. Algeria.
Resumo
Compreender a influência das variações de temperatura na germinação de sementes de trigo duro é de importância prática para agricultores, pesquisadores e tomadores de decisões agrícolas. A capacidade das sementes de germinar de forma ideal é crucial para garantir rendimentos satisfatórios das colheitas. O principal objetivo deste trabalho é analisar o impacto das variações de temperatura na viabilidade e vigor das sementes de trigo duro na Argélia. Para atingir esse objetivo, adotamos uma abordagem abrangente que se concentrou principalmente no estudo dos parâmetros morfológicos, fisiológicos, bioquímicos e tecnológicos (como germinação, cinética de crescimento de mudas, proteína solúvel, prolina e teor de açúcar solúvel) das sementes. De fato, ao longo de um período de dois anos, quatro variedades de trigo (Bousselam, Setifis, Megress, Saoura) foram submetidas a duas condições distintas de temperatura e umidade durante o estudo. A primeira condição envolveu temperaturas ambientes flutuando entre 11°C e 37°C, com umidade relativa variando de 30% a 85%. Entretanto, a segunda condição envolveu o armazenamento das sementes em uma câmara fria a uma temperatura de -04°C e umidade de 12%. Os resultados mostram variações significativas na taxa de germinação de todas as variedades sob ambas as condições de temperatura F=40, 54 P=0,000, com a variedade Bousselam (B) se destacando como a mais resistente às mudanças térmicas. De acordo com os resultados obtidos, a temperatura afeta o crescimento das raízes da variedade S, enquanto as variedades Bousselam (B), Megress(M), Saoura (SW) não são afetadas; além disso, o armazenamento sob ambas as condições de temperatura tem impacto no número médio de raízes e no comprimento dos coleóptilos. As análises de proteínas revelaram efeitos significativos do armazenamento sob condições ambientais (F=4,70 P=0,000), enquanto na câmara fria, esses efeitos foram atenuados. Além disso, o armazenamento mostrou variações no teor de açúcar solúvel e prolina, com diferentes respostas sob as duas condições de temperatura. Esses resultados ressaltam a importância de entender as respostas de variedades de trigo duro a condições térmicas variáveis, oferecendo perspectivas úteis para melhorar práticas agrícolas e preservar a qualidade das sementes em ambientes em mudança.
Palavras-chave: Temperatura. Germinação. Trigo duro. Sementes. Argélia.
Introduction
The durum wheat is one of the most important crops economically and nutritionally (ZINGALE et al., 2019).
Seed quality is affected by physiological parameters and sanitary conditions (AYA et al., 2011) and the germination is a vital process in the plant life cycle, marking the beginning of the growth of a new plant from a seed (RAJJOU et al., 2012).
Temperature is one important factor triggering seed germination (CATARA et al., 2015). Emerging evidence suggests that plants perceive the fluctuating temperatures at multiple levels and climate change and extreme temperatures decrease global crop yields (DING et al., 2020).
Some findings suggest that temperature-mediated epigenetic regulation is promising in plants during germination (WANG et al, 2024) and the molecular mechanisms of integration of light and temperature information are diverse (PANERO et al., 2024).
It was observed that the increase in the average maximum temperature positively influenced the number of falls, in the period corresponding to the end of grain filling and during physiological maturation; on the other hand, it was observed that the average minimum temperature negatively affected the number of falls during grain filling and physiological maturation (GUARIENTI et al., 2024).
The link between optimizing agricultural practices and the phenomenon of seed germination is crucial for ensuring the success of agricultural production. Agricultural practices encompass a wide range of activities, including soil preparation, seed selection, planting techniques, irrigation, fertilization, pest and disease management, and harvesting. These practices directly influence the conditions under which seeds germinate, as well as the subsequent growth and development of plants.
Through this study, we aim to contribute to a better understanding of the germination mechanisms of durum wheat and the impact of temperature variations on this process. This knowledge could lead to more effective agricultural practices and better management of durum wheat crops, thereby contributing to food security and environmental sustainability.
Material and methods
Plant material
Our experience was carried out at the Plant Improvement Laboratory in Badji Mokhtar Annaba in Algeria. The basic plant material comprised four durum wheat varieties, namely Bousellam (Pedigree: Heider/Martes/Huevos de Oro. ICD-414; CIMMYT-ICARDA), Setifis, Megress (Pedigree, Ofanto/Waha//MBB, ITGC, ARS, Setif, Algeria) and Saoura. The wheat seed was kindly provided by the ITGC station (Institut Technique des Grandes Cultures – Algeria) (Table 1).
Table 1 – The four durum wheat varieties – Algeria.
|
|
Pedigree |
Origin |
Bousellam |
Heider/Martes/Huevos de Oro. ICD-414 |
CIMMYT-ICARDA |
Setifis |
Bousselam/Ofanto |
ITGC-Setif |
Megress |
Ofanto/Waha//MBB, ITGC, ARS |
Setif, Algeria |
Saoura |
Ofanto/waha |
Saoura |
Experimental setup and experiment conditions
It was conducted according to a completely randomized protocol with three replicates (R1, R2, R3) and three distinct treatments (Figure 1):
- Treatment T (control): use of fresh durum wheat seed;
- Favorable conditions: treatment (cold room): exposure of durum wheat seeds to specific temperature conditions, with a temperature of -04°C and a humidity of 12% in a cold room;
- Unfavorable conditions: treatment (laboratory): storage of durum wheat seeds under conditions of ambient temperature (between 10 and 36°C) and humidity (between 48-83%) in the laboratory.
Figure 1 – Experimental setup for germination and vigor test (C: control, FC: favorable conditions, UFC: unfavorable conditions, R: repetition).
These conditions were set up to assess the effects of different temperatures on durum wheat seed germination.
The disinfected seeds were placed on water-soaked cotton wool in Petri dishes. The leaves were harvested after 14 days for analysis of various parameters.
Standard germination test
To determine the germination percentage, random seed samples were tested by subjecting them to favorable germination conditions. The average repeating number is 100 seeds for each treatment (25 seeds per Petri dish) (Figure 2).
Figure 2 – Germination test.
The percentage is calculated after 3 days of germination formatting by the following formula:
Germination (%) = Number of germed seeds x 100 / Number of seeds to be germinated
Coleoptile and root length
The length of the coleoptile and the root of each seedling resulting from germination is measured with a measured with a graduated ruler for each variety and each treatment. The measurements of this parameter are made after 14 days of germination. The number of roots by plantlet was also counted.
Soluble sugar content
The determination of soluble sugars was carried out using the method described by Schields and Burnett (1960), also known as the anthrone method in sulfuric acid medium.
Proline content
Proline, or pyrrolidine-2-carboxylic acid, is one of the twenty major amino acids that constitute proteins. Proline is easily oxidized by ninhydrin or triketohydrindene. It will be determined according to the method of Troll and Lindsley (1995), modified by Monneveux and Nemmar (1986).
Total protein content
Total protein content is measured using the Bradford method (1976) with bovine serum albumin (B.S.A.) as the standard.
Statistical data analysis
The statistical analysis of the data was conducted using Tukey’s test and two-way analysis of variance (ANOVA), using the specific data analysis and processing software MINITAB version 15.1, to determine significant differences between the means of homogeneous groups. The differences are displayed as follows:
Significant when *p < 0.05;
Highly significant when ** p < 0.01;
Very highly significant when *** p < 0.001.
Results
Germination test
After a four-day germination period, the germination percentages of seeds under two temperature conditions were observed for four durum wheat varieties, as shown in the Figure 3. The germination rate of untreated seedsSeeds without treatment.) ranged from 90% to 95%, for Saoura and Setifis, respectively.
Figure 3 – Effect of temperature variations on the germination capacity of four durum wheat varieties, subjected to storage after three days of soaking.
The figure shows that, whatever the variety, the germination capacity of stressed eeds are reduced compared with the control under both temperature conditions.
It should be noted that variety B is the most resistant and showed a germination rate of 94% under cold room conditions and 88.33% under room temperature conditions.
It is important to note that, under high temperature conditions, variety SW stands out by presenting a lower germination rate than the control, with an average of 67.66%. The analysis of variance reveals a very highly significant difference, both for the variety factor (F=40, 54 P=0,000); these results are consistent for all varieties.
Root length
The results show that variations in temperature have an impact on root growth in variety S, while varieties B, M and SW do not seem to be affected, as shown in the Figure 2.
Root growth of control seeds varied between 100.16 mm and 150.66 mm for Saoura (SW) and Setifis (S), respectively. The high temperatures affected root growth of seeds of variety S, which fell from 150.66 mm in the control to 117.5 mm.
Although varieties B, M and SW showed no impact under the influence of temperature conditions, variety S reacted in a similar way under cold temperature conditions, with a reduction in root growth of up to 129.16 mm compared with the control (Figure 4). Analysis of variance, taking into account temperature and variety factors, did not reveal any significant differences. (F=0, 14 P=0,934).
Figure 4 – Effect of temperature variations on root growth of four durum wheat varieties.
Coleoptiles length
Unlike root growth, coleoptiles growth is negatively affected by seed storage under both ambient and cold storage conditions (Figure 5).
Coleoptiles growth in control seeds ranged from 115.16 mm to 156.83 mm for SW and B, respectively. A decrease in coleoptiles length was observed under lower temperature conditions for the three varieties B, S and M (120), ranging from 64 mm to 144 mm for B and S, respectively. The length of the coleoptiles of the seeds of the four varieties studied was also more affected under high temperature conditions than under lower temperature conditions, varying from 88.16 mm to 100.33 mm in S and SW varieties, respectively. Analysis of variance gives a highly significant difference for the treatments (F=1, 45 P=0,236).
Figure 5 – Effect of temperature variations on the growth of coleoptiles of four durum wheat varieties.
Average root number
Storage under favorable and unfavorable temperature conditions had an impact on the average number of roots, as shown in the Figure 6. Root growth in control seeds varied between 24.66 mm and 48.5 mm for SW and M, respectively.
A decrease in the average number of roots was observed under lower temperature conditions (-04°C) linked to storage in the group of varieties S(30cm), M and SW, rising from 18.66 mm to 35 mm for SW and M, respectively, while the variety B was not affected.
The average number of roots of the four varieties studied after storage under high temperature conditions (between 10 and 36°C) was also more affected than under cold temperature conditions, varying from 21.66 mm to 31.83 mm in varieties and B, respectively. Despite this, analysis of variance did not reveal any significant differences (F=3, 01 P=0,037).
Figure 6 – Effect of temperature variations on the average number of roots of four durum wheat varieties.
Total protein content
In the control group, an average accumulation of total protein of 13.56 to 20.64 μg/100 mg fresh matter was observed in the two varieties S and M, respectively (Figure 7).
A significant influence of storage was observed in all four varieties (B, S, M, SW), which showed considerably lower values than the control under high temperature conditions, with a minimum of 10.92 μg/100 mg recorded in the B variety and a maximum of 11.28 μg/100 mg noted in the SW variety. On the other hand, there was no influence under low temperature conditions, 21 μg/100 mg and 19 μg/100 mg was observed in the two varieties B and SW, respectively.
Analysis of variance for the temperature level factor revealed a highly significant difference in all sources of variation (F=14, 70 P=0,000); these results are also consistent for the variety factor and the interaction factor (variety × T level).
Figure 7 – Effect of temperature variations on the total protein content of four durum wheat varieties.
Soluble sugar content
In the control group, an average accumulation of soluble sugars ranging from 5.69 to 8.96 μg/100 mg fresh matter was observed in the two varieties SW and B, respectively (Figure 8).
A significant influence on germination conditions was noted in three varieties (B, S, SW), which showed high values compared with the control under favorable conditions. A minimum of 6.03 μg/100 mg ecorded in the variety SW and a maximum of 13.18 μg/100 mg oted in the variety B;8 μg/100 mg for the S variety.
Variety M, on the other hand, showed a decrease in soluble sugars to 5.53 μg/100 mg. In conditions of high temperature conditions, the soluble sugar content increases more than in favorable conditions, with a minimum of 9.77 μg/100 mg recorded in variety M and a maximum of 16.68 μg/100 mg noted in variety B.
Analysis of variance for the temperature level factor (treatment) and varieties reveals a highly significant difference (F=43, 91 P=0,000).
Figure 8 – Effect of temperature variations on the soluble sugar content of four durum wheat varieties.
Proline content variation
Under untreated conditions, an average proline accumulation of 16.76 to 28.08 μg/100 mg fresh matter was observed in the two varieties M and B, respectively (Figure 9).
A notable effect of germination conditions was noted in all four varieties (B, S, M, SW), with high values compared with the control under conditions of lower temperature, with a minimum of 27.22 μg/100 mg recorded in the M variety and a maximum of 36.06 ug/100 mg recorded in the S variety.32 ug/100 mg for B variety and 33 ug/100 mg to SW variety.
Under conditions of high temperature, the proline content increases more than under low temperature conditions, with a minimum of 35.14 ug/100 mg in variety B and a maximum of 54 ug/100 mg in variety SW.
Data variance analysis revealed a highly significant difference in all sources of variation, (F=44, 52 P=0,000).
Figure 9 – Effect of temperature variations on the proline content of four durum wheat varieties.
Discussion
The seeds enter in a period of rapid decline during which some seeds fail to germinate completely, while others germinate and grow normally (ABDUL-BAKI and ANDERSON, 1972). It is for this reason that viability tests, using different seed vigour tests, are very important, since vigour tests often give results that correlate better with germination results in the field, under unfavourable environmental conditions, than results obtained by applying the standard germination test in the laboratory (JOHNSON and WAX, 1978).
Germination is defined as the appearance and development, from the seed embryo, of those essential organs which are indicative of the seed’s ability to produce a normal plant under favourable conditions (JUSTICE, 1972). The germination rate is expressed by the percentage of pure seeds that produce normal seedlings or by the number of germinated seeds per unit weight of the sample (WILLAN, 1992).
According to the results obtained during our study, we observed a decrease in the germination rate as a function of the duration and temperature conditions applied for all the varieties studied.
We found that seeds stored under ambient temperature conditions were the most affected, with lower germination rates than seeds stored under low-temperature conditions, according to Multon (1982), storage time is a factor that amplifies deterioration phenomena. Booth and Sowa (2001) and Srivastava (2002) believe that the weaker germination of aged seeds is due to the natural ageing process; even when stored under controlled temperature and humidity conditions, seeds gradually lose their viability.
In fact, storage can affect seed viability and vigour, depending on time and various conditions (PANOBIANCO et al., 2007). Cereal seeds, including wheat, are in most cases stored under normal storage conditions and as these conditions vary according to the season of the year, seeds stored in cloth or paper bags are tempered and easily exchange moisture with the ambient air (COPELAND and McDONALD, 2001; VOLENIK et al., 2006).
If the moisture balance between the seeds and the ambient air leads to an increase in the humidity of the seeds, the deterioration process increases with the concomitant temperature inside the seeds, leading to a reduction in germination and vigour. Similarly, seeds with low moisture content stored in conditions unfavourable to the acquisition of additional moisture usually maintain high germination (EDMOND, 1962; KEARNS and TOOLE, 1939; TOOLE et al., 1948).
In addition, Wallace and Sinha (1962) reported a negative correlation between storage temperature and germination. Our results are in line with numerous studies that report a reduction in germination capacity after long periods of storage under high temperature conditions (GOVENDER et al., 2008). Other studies have shown that germination is negatively correlated with temperature and storage period in many species such as Phaseolus vulgaris (RANI et al., 2013), Hordeum vulgare and Avena sativa L. (WHITE et al., 1999), Triticum durum (KARUNAKARAN et al., 2001; NITHYA et al., 2011) and Secale cereale L. (SATHYA et al., 2008, 2009).
Nobbe (1876), recognised that individual seed properties, such as germination rate and seedling growth, varied within the same seed lot and that seed lot averages also varied very often. This reduction in germination during storage may also be due to visible and non-visible fungal microflora, which can damage the embryo (CHRISTENSEN, 1969) and deplete nutrient reserves (CHRISTENSEN, 1973) through the production of toxic metabolites (HARMAN and NASH, 1972; LACEY, 1975; BHATTACHARYA and RAHA, 2002; CAID et al., 2008).
Measuring plant growth can be considered an excellent, inexpensive and reliable test for estimating seed vigour. Our results show a reduction in root and leaf elongation of wheat seedlings with the appearance of abnormal seedlings as a function of temperature in all the varieties studied.
Generally, when the germination percentage of a seed lot decreases, many of the seedlings obtained are abnormal (TOOLE et al., 1948; ABDUL-BAKI and ANDERSON, 1972), these seedlings may have poorly developed roots and/or stems (TOOLE et al., 1948) and have necrotic root meristems (ABDUL-BAKI and ANDERSON, 1972).
According to Coin et al. (1995) storage in different conditions leads to a reduction in vigour and a slowdown in growth, resulting in a loss of germination capacity. Deteriorated seeds, if they germinate at all, often produce slow-growing seedlings (ABDUL-BAKI and ANDERSON, 1972).
The drop in protein content, along with the increase in proline and soluble sugar content in the four varieties studied is part of the overall biochemical degradation that grains undergo during storage. These same changes have been observed in pea seeds (KALPANA and RAO, 1994) and wheat seeds artificially aged (DELL AQUILA, 1994).
The degradation of soluble proteins is thought to affect gliadins and glutenins in particular, whose role in bread-making quality is well established. The increase in proline content during accelerated ageing could be explained as a consequence of protein degradation or simply as a response to the heat stress (40°C) to which the seeds are subjected during processing.
The increase in reducing sugars in the four wheat varieties studied is thought to be the result of increased starch degradation during treatment. These sugars are thought to initiate Amadori and Maillard reactions (CALUCCI et al., 2004; SUN and LEOPOLD, 1995), which are responsible for grain browning and, above all, are associated with loss of viability during storage.
The decrease in protein content observed in stored seeds may be the result of the formation of free radicals which denature, oxidise or degrade these proteins to form carbonyl derivatives (SHACTER et al., 1994; ALAYAT, 2015).
Proline was quantified in the leaves of wheat seedlings. The results showed a very significant increase in prolin levels in the leaves of seedlings from old seed in all the varieties studied. Proline is a free amino acid considered to be a bioindicator of stress. These molecules are solutes of cellular metabolism that protect plants against various abiotic stresses, through osmotic readjustment, which maintains cellular turgidity and water uptake under hyperosmotic conditions (BENKADOUR, 2014).
This accumulation of proline is due to a disturbance in metabolism and/or a nitrogen storage process necessary for cell survival (SIVARAMAKRISHNAN et al., 1988; BENKADOUR, 2014).
The identification of a clear correlation between high temperatures and reduced viability underlines the crucial importance of careful temperature management during seed storage. Excessively high temperatures can cause irreversible damage to the cellular structures and biological mechanisms of seeds.
The reduction in viability observed at high temperatures can have a direct impact on the germination rate. This relationship highlights the need for specific measures to be taken to maintain optimum storage conditions in order to guarantee an optimal germination rate.
Variability between wheat genotypes can influence the way they react to high temperatures. Some genotypes may be more resilient to these conditions, emphasising the importance of selecting suitable varieties to ensure that seed viability is preserved.
The results of this study have direct implications for agricultural practices, particularly seed storage. The implementation of appropriate storage practices, based on a detailed understanding of seed responses to temperature, can help to guarantee the quality and viability of seeds used for crops.
Conclusion
In conclusion, the results obtained underline the significant influence of temperature on the characteristics of wheat seeds, highlighting several key observations. This research revealed a significant correlation between temperature conditions and the germination rate of wheat seeds. It has been observed that high temperatures are associated with a reduction in seed viability, highlighting the importance of meticulous management of thermal conditions during storage. Variations in temperature had specific repercussions on root and coleoptiles growth.
Biochemical analyses revealed substantial variations in the accumulation of total proteins, soluble sugars and proline in response to changes in temperature. These results suggest a metabolic plasticity of wheat seeds in the face of thermal stress. The differences observed between genotypes highlight the importance of selecting wheat varieties adapted to specific thermal conditions. This genetic adaptability could play a key role in maintaining crop productivity in the face of climatic fluctuations.
In conclusion, the results discussed highlight the crucial importance of managing temperature conditions to ensure the viability of wheat seed. These findings have direct implications for farming practices and call for a proactive approach to seed storage management in order to preserve the genetic quality and germination performance of wheat seed.
Interest conflicts
There was no conflict of interest of the authors.
Authors’ contribution
Lina Manel Djendi and Amel Soussa – original idea, field work, reading and interpretation of works and writing; Nabila Bouhaddouda and Amina Dridi – writing and corrections; Zeriri Ibtissem and Hiba Daas – guidance, corrections and revision of the text.
References
ABDUL-BAKI, A. A.; ANDERSON, J. D. Physiological and biochemical deterioration of seeds. Seed Biology, v. 2, p. 283-315, 1972. https://doi.org/10.1016/B978-0-12-424303-3.50010-5
ALAYAT, A. Etude de l’Impact Toxicologique de Certains Agents Chimiques sur la Qualité des Céréales: cas du Blé et de l’Orge. 231p. Thèse (Doctorat) – Université Annaba, Algerie, 2015.
AYA, A. N. N.; VROH-BI, I.; KOUAMÉ, P. L.; ZORO BI, I. A. Bases génétiques et biochimiques de la capacité germinative des graines: implications pour les systèmes semenciers et la production alimentair. Sciences & Nature, v. 8, n. 1, p. 119-137, 2011. https://www.ajol.info/index.php/scinat/article/view/92849
BENKADDOUR, M. Modifications physiologiques chez des plantes de blé (Triticum durum Desf) exposées à un stress salin. 101p. Thèse (Doctorat) – Université Annaba, Algerie, 2014.
BHATTACHARYA, K.; RAHA, S. Deteriorative changes of maize, groundnut and soybean seeds by fungi in storage. Mycopathologia, v. 155, p. 135-141, 2002. https://doi.org/10.1023/A:1020475411125
BOOTH, D. T. ; SOWA, S. Respiration in dormant and non-dormant bitterbrush seeds. Journal of Arid Environments, v. 48, p. 35-39, 2001. https://doi.org/10.1006/jare.2000.0737
BRADFORD, M. M. A rapid and sensitive method for the quantitation of microgram quantities of protein utilizing the principle of protein-dye binding. Analytical Biochemistry, v. 72, v. 1-2, p. 248-254, 1976. https://doi.org/10.1016/0003-2697(76)90527-3
CAID, H. S.; ECCHEMMAKH, T.; ELAMRANI, A.; KHALID, A.; BOUKROUTE, A.; MIHAMOU, A.; DEMANDRE, C. Altérations accompagnant le vieillissement accéléré de blé tendre. Cahiers Agricultures, v. 17, n. 1, p. 39-44, 2008. https://doi.org/10.1684/agr.2008.0157
CALUCCI, L.; CAPOCCHI, A.; GALLESCHI, L.; GHIRINGHELLI, S.; PINZINO, C.; SAVIOZZI, F.; ZANDOMENEGHI, M. Antioxydants, free radicals, storage proteins, puroindolines and proteolytic activities in bread wheat (Triticum aestivum) seeds during accelerated aging. Journal of Agricultural and Food Chemistry, v. 52, n. 13, p. 4274-4281, 2004. https://doi.org/10.1021/jf0353741
CATARA, S.; CRISTAUDO, A.; GUALTIERI, A.; GALESI, R.; IMPELLUSO, C.; ONOFRI, A. Threshold temperatures for seed germination in nine species of Verbascum (Scrophulariaceae). Seed Science Research, v. 26, n. 1, p. 30-46, 2016. https://doi.org/10.1017/S0960258515000343
CHRISTENSEN, C. M. Loss of viability in storage: microflora. Seed Science and Technology, v. 1, n. 3, p. 547-562, 1973.
CHRISTENSEN, C. M.; KAUFMANN, H. H. Grain Storage: The Role of Fungi in Quality Loss. University of Minnesota: Academic Press, 1969, 172p.
COIN, M.; ANDERSON, J.; MILLER, T. The impact of storage conditions on seed vigour and germination capacity. Seed Science and Technology, v. 23, n. 4, p. 305-315, 1995.
COPELAND, L. O.; McDONALD, M. B. Principles of Seed Science and Technology. New York, NY: Springer, 2001, 467p. https://doi.org/10.1007/978-1-4615-1619-4
DELL AQUILA, A. Wheat seed ageing and embryo protein degradation. Seed Science Research, v. 4, n. 3, p. 293-298, 1994. https://doi.org/10.1017/S0960258500002324
DING, Y.; SHI, Y.; YANG, S. Molecular regulation of plant responses to environmental temperatures. Molecular Plant, v. 13, n. 4, p. 544-564, 2020. https://doi.org/10.1016/j.molp.2020.02.004
EDMOND, J. B. Studies of the storage of vegetable seed under warm-humid conditions. Mississippi State University, Agricultural Experiment Station, State College, Miss., 1962.
GOVENDER, V.; AVELING, T. A. S.; KRITZINGER, Q. The effect of traditional storage methods on germination and vigour of maize (Zea mays L.) from northern KwaZulu-Natal and southern Mozambique. South African Journal of Botany, v. 74, n. 2, p. 190-196, 2008. https://doi.org/10.1016/j.sajb.2007.10.006
GUARIENTI, E. M.; CIACCO, C. F.; CUNHA, G. R.; DEL DUCA, L. J. A.; CAMARGO, C. M. O. nfluence of minimum and maximum temperature in wheat industrial quality characteristics and in grain yield. Food Science and Technology, v. 24, n. 4, p. 505-515, 2004. https://doi.org/10.1590/S0101-20612004000400005
HARMAN, G. E.; NASH, G. Deterioration of stored pea seed by Aspergillus ruber: evidence for the involvement of a toxin. Phytopathology, v. 62, n. 2, p. 209-212, 1972. https://www.apsnet.org/publications/phytopathology/backissues/Documents/1972Abstracts/Phyto62_209.htm
JOHNSON, R. R.; WAX, L. M. Relationship of soybean germination and vigour test to field performance. Agronomy Journal, v. 70, n. 2, p. 273-278, 1978. https://doi.org/10.2134/agronj1978.00021962007000020014x
JUSTICE, O. L. Essentials of seed testing. In: KOZLOWSKI, T. T. Seed Biology. Volume 3. Universidade de Minnesota: Academic Press, 1972, 422p.
KALPANA, R.; RAO, K. V. M. Absence of the role of lipid peroxidation during accelerated ageing of seeds of pigeonpea (Cajanus cajan (L.) Millsp.) cultivars. Seed Science and Technology, v. 22, n. 2, p. 253-260, 1994.
KARUNAKARAN, C.; MUIR, W. E.; JAYAS, D. S.; WHITE, N. D. G.; ABRAMSON, D. Safestorage time of high moisture wheat. Journal of Stored Products Research, v. 37, n. 3, p. 303-312, 2001. https://doi.org/10.1016/S0022-474X(00)00033-3
KEARNS, V; TOOLE, E. H. Relation of temperature and moisture content to longevity of Chewings fescue seeds. Technical Bulletins 670, United States Department of Agriculture, 1939, 27p. https://doi.org/10.22004/ag.econ.167929
LACEY, J. Molding of grain in relation to mycotoxin formation. International Journal of Environmental Studies, v. 8, n. 1-4, p. 183-186, 1975. https://doi.org/10.1080/00207237508709731
MONNEVEUX, P.; NEMMAR, M. Contribution à l’étude de la résistance à la sécheresse chez le blé tendre (Triticum aestivum L.) et chez le blé dur (Triticum durum Desf.): étude de l’accumulation de la proline au cours du cycle de développement. Agronomie, v. 6, n. 6, p. 583-590, 1986. https://doi.org/10.1051/agro:19860611
MULTON, J. L. Conservation et stockage des grains et graines et produits dérivés: céréales, oléagineaux, protéagineux, aliments pour animaux. Paris: Lavoisier – Technique et Documentation, 1982.
NITHYA, U.; CHELLADURAI, V.; JAYAS, D. S.; WHITE, N. D. G. Safe storage guidelines for durum wheat. Journal of Stored Products Research, v. 47, n. 4, p. 328-333, 2011. https://doi.org/10.1016/j.jspr.2011.05.005
NOBBE, F. Handbuch der Samenkunde. Berlin: Verlag von Wiegandt, Hempel und Parey, 1876.
PANERO, I.; FIORENTTINO, F.; LA MONTAGNA, D.; CROCENZI, G.; ATTORRE, F.; FABRINI, G. Germination ecology of Phytolacca americana L. in its invasive range. Plant Species Biology, early view, 2024. https://doi.org/10.1111/1442-1984.12483
PANOBIANCO, M.; VIEIRA, R. D.; PERECIN, D. Electrical conductivity as an indicator ofpea seed aging of stored at different temperatures. Scientia Agricola, v. 64, n. 2, p. 119-124, 2007. https://doi.org/10.1590/S0103-90162007000200003
RAJJOU, L.; DUVAL, M.; GALLARDO, K.; CATUSSE, J.; BALLY, J.; JOB, C.; JOB, D. Seed germination and vigor. Annual Review of Plant Biology, v. 63, p. 507-533, 2012. https://doi.org/10.1146/annurev-arplant-042811-105550
RANI, P. R.; CHELLADURAI, V.; JAYAS, D. S.; WHITE, N. D. G.; KAVITHA-ABIRAMI, C. V. Storage studies on pinto beans under different moisture contents and temperature regimes. Journal of Stored Products Research, v. 52, p. 78-85, 2013. https://doi.org/10.1016/j.jspr.2012.11.003
SATHYA, G.; JAYAS, D. S.; WHITE, N. D. G. Safe storage guidelines for rye. Canadian Biosystems Engineering, v. 50, p. 3.1-3.8, 2008. https://library.csbe-scgab.ca/all-publications/2617:safe-storage-guidelines-for-rye
SATHYA, G.; JAYAS, D. S.; WHITE, N. D. G. Safe storage guidelines for canola as the seeds slowly dry. Canadian Biosystems Engineering, v. 51, p. 3.29-3.38, 2009. https://library.csbe-scgab.ca/all-publications/2646:safe-storage-guidelines-for-canola-as-the-seeds-slowly-dry
SCHIELDS, R.; BURNETT, W. Determination of protein-bound carbohydrate in serum by modified anthrone method. Analytical Chemistry, v. 32, n. 7, p. 885-886, 1960. https://doi.org/10.1021/ac60163a053
SHACTER, E.; WILLIAMS, J. A.; LIM, M.; LEVINE, R. L. Differential susceptibility of plasma proteins to oxidative modification: examination by western blot immunoassay. Free Radical Biology and Medicine, v. 17, n. 5, p. 429-437, 1994. https://doi.org/10.1016/0891-5849(94)90169-4
SIVARAMAKRIAHNAN, S.; VILLOO, Z.; PATELL, D. J.; FLOWER, J. M. Peacock Proline accumulation and nitrate reductase activity in contrasting sorghum lines during mid-season drought stress. Physiologia Plantarum, v. 74, n. 3, p. 418-426, 1988. https://doi.org/10.1111/j.1399-3054.1988.tb01997.x
SRIVASTAVA, L. M. Plant Growth and Development. Hormones and the Environment. Oxford, Academic Press, 2002, 722p.
SUN, W. Q., LEOPOLD, A. C. The Maillard reaction and oxidative stress during aging of soybean seeds. Physiologia Plantarum, v. 94, n. 1, p. 94-105, 1995. https://doi.org/10.1111/j.1399-3054.1995.tb00789.x
TOOLE, E.; TOOLE, H.; GORMAN, E. A. Vegetable-seed storage as affected by temperature and relative humidity. Technical Bulletins 972, United States Department of Agriculture, 1948, 24p. http://doi.org/10.22004/ag.econ.170310
TROLL, W.; LINDSEY, J. A photometricmethod for the determination of proline. Journal of Biological Chemistry, v. 215, n. 2, p. 655-660, 1955. https://pubmed.ncbi.nlm.nih.gov/13242563/
VOLENIK, M.; ROZMAN, V.; KALINOVIC, I.; LISKA, A.; KIS, D.; SIMIC, B. Influence of relative humidity and temperature on changes in grain moisture in stored wheat and sunflower. In: 9th International Working Conference on Stored Product Protection. Proceedings… Campinas, Brazil, p. 24-29, 2006. https://www.researchgate.net/publication/Influence_of_relative_humidity_and_temperature_on_changes_in_grain_moisture_in_stored_wheat_and_sunflower.pdf
WALLACE, H. A. H.; SINHA, R. N. Fungi associated with hot spots in farm stored grain. Canadian Journal of Plant Science, v. 42, p. 130-141, 1962. https://doi.org/10.4141/cjps62-016
WANG, Y.; SUN, X.; PENG, J.; LI, F.; ALI, F.; WANG, Z. Regulation of seed germination: ROS, epigenetic, and hormonal aspects. Journal of Advanced Research, In Press, 2024. https://doi.org/10.1016/j.jare.2024.06.001
WHITE, N. D. G.; HULASARE, R. B.; JAYAS, D. S. Effects of storage conditions on quality loss of hull-les and hulled oats and barley. Canadian Journal of Plant Science, v. 79, n. 4, p. 475-482, 1999. https://doi.org/10.4141/P98-115
WILLAN, A. Seed germination rates: measurement and significance. Seed Technology Journal, v. 15, n. 2, p. 99-110, 1992.
ZINGALE, S.; SPINA, A.; INGRAO, C.; FALLICO, B.; TIMPANARO, G.; ANASTASI, U.; GUARNACCIA, P. Factors affecting the nutritional, health, and technological quality of durum wheat for pasta-making: a systematic literature review. Plants, v. 12, n. 3, p. 1-37, 2023. https://doi.org/10.3390/plants12030530
Received on March 3, 2024
Returned for adjustments on August 2, 2024
Received with adjustments on August 20, 2024
Accepted on August 28, 2024