Variations in grain textural and starch physicochemical properties of waxy maize across contrasting ecological regions

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Variations in these natural ecological conditions can result in differences in quality within the same variety grown in different regions. The grain texture and starch physicochemical properties were examined across five different sites in Hebei Province, including Hengshui (HS), Baoding (BD), Tangshan (TS), Qinhuangdao (QHD), and Chengde (CD). Significant divergence of grain quality existed across the five sites. The grain hardness was highest at the TS site, while the adhesiveness was optimal at the HS site. The BD site exhibited the highest soluble sugar content, and the TS site had the highest total starch content. Notably, the amylopectin content at the QHD site was significantly higher than at the other locations. Regarding starch structure, the starch granules from the TS site were the largest and demonstrated the highest relative crystallinity. In contrast, the starch granules from the HS site were smaller and more uniformly distributed. Analysis of starch functional properties revealed that the peak viscosity of starch was significantly higher at BD and TS sites. The enthalpy value and swelling power of starch were optimal at the TS site, whereas the HS site exhibited higher starch light transmittance but weaker stability. Further analysis indicated that these differences were significantly associated with the soil conditions and climatic factors at each site. Overall, this study provide a theoretical basis for the application of waxy maize starch in the food industry and for improving starch quality through breeding. Biological sciences/Biochemistry Biological sciences/Plant sciences Waxy maize starch ecological region Figures Figure 1 Figure 2 Figure 3 Introduction Waxy maize is a distinctive type of maize extensively cultivated in China, its starch is almost entirely composed of amylopectin, which provides high viscosity, a low retrogradation rate, and good stability [ 1 ]. These unique physicochemical characteristics of the starch give waxy maize a high value for processing and utilization, resulting in its extensive application in the food industry and in industrial products, including thickeners, emulsifiers, and adhesives [ 2 ]. As the primary component of waxy maize, starch is a key determinant of its quality. Specifically, the physicochemical properties of the starch are directly influenced by the ratio and structure of amylose to amylopectin, which, in turn, determine its suitability for specific applications [ 3 – 5 ]. Starch possesses a multi-scale hierarchical structure, characterized by features, including amylopectin chain length, particle morphology, crystalline structure, and molecular conformation. Key indicators for evaluating waxy corn starch properties include amylopectin content, particle size distribution, crystallinity, and gelatinization temperature. Notably, these characteristics are susceptible to environmental and cultivation factors during the grain formation period, including temperature, waterlogging stress at the flowering stage, and nitrogen level [ 6 – 7 ]. Furthermore, exogenous substances, including coronatine and salicylic acid, can improve waxy corn grain yield and starch quality by regulating the activity of key enzymes involved in starch synthesis and the expression of related genes [ 8 – 9 ]. The growth environment is a critical regulatory factor in starch macromolecule biosynthesis. Studies indicate that temperature fluctuations during the grain-filling stage can affect the activities of starch synthase and branching enzymes, thereby altering the amylopectin chain-length distribution and crystal packing modes [ 9 – 11 ]. Drought stress reduces the starch content in waxy maize, increases the levels of total protein, globulin, and glutelin, and results in an increase in starch average particle size, amylopectin average chain length, and relative crystallinity. Furthermore, the impact of drought stress is more pronounced during the kernel-formation stage than during the grain-filling and enrichment stages [ 12 ]. Variations in the ecological region's environmental conditions contribute to changes in starch properties. A study comparing broomcorn millet starch from Yangling and Yulin in Shaanxi Province observed that in the warmer Yangling region, the starch exhibited a higher proportion of long amylopectin chains and greater relative crystallinity. This was accompanied by an increase in gelatinization temperature, increased shear resistance, and a decrease in retrogradation value [ 13 ]. Furthermore, a study on waxy maize has confirmed that high-temperature stress during kernel formation increases the average particle size of starch granules and the proportion of long amylopectin chains, thereby altering their gelatinization and thermodynamic properties [ 14 ]. A comparative study of heat-sensitive and heat-tolerant waxy corn varieties revealed that high temperatures during grain development result in surface depressions on starch granules, increased particle size, a higher proportion of long amylopectin chains, and elevated relative crystallinity. Subsequently, these changes reduce the gelatinization viscosity and enthalpy of starch while increasing the retrogradation rate [ 15 ]. Historically, waxy maize was considered a specialty crop. However, due to its unique starch composition, strong processing adaptability, and outstanding nutritional value, it has now emerged as a potential crop for diversifying food industry raw materials and increasing the added value of agricultural products. Waxy maize is extensively cultivated across China, spanning multiple ecological regions from the humid, hot south to the arid north. These regions exhibit significant differences in temperature, precipitation, and soil fertility. Variations in these natural ecological conditions can result in differences in quality within the same variety grown in different regions. Additionally, the extent of these differences can be comparable to that observed among different varieties within the same area [ 16 – 17 ]. Therefore, this study hypothesizes that waxy maize produced across different ecological regions can exhibit significant variation in grain texture characteristics and in the physicochemical properties of starch. These differences would directly affect the potential of waxy maize in both food and non-food industries, as the development of specific products depends on starch raw materials with the corresponding properties. This study used the novel fresh-eating waxy maize variety "Sweet Waxy" and its representative cultivar Jingkenuo 768 [ 18 ] as experimental materials. These materials were cultivated across four ecological regions in Hebei Province, characterized by contrasting ecological factors. This study aimed to identify patterns of grain quality variation in waxy maize across these regions by analyzing the relationships among grain texture characteristics, starch physicochemical properties, and environmental conditions in different ecological regions of Hebei Province. The findings can provide theoretical and technical references for high-quality waxy maize cultivation in the province. The specific objectives of this study were as follows: (1) analyzing the grain texture, starch particle morphology, crystalline and infrared spectral structural features of waxy maize from different ecological regions in Hebei Province and evaluating the functional properties of its starch; (2) investigating the association mechanisms between the grain texture of waxy maize and the structural and functional properties of its starch; (3) identifying the influences of meteorological ecological factors and soil conditions on the grain texture and physicochemical properties of starch in waxy maize. Materials and Methods Experimental site conditions Five experimental sites were selected within Hebei Province, including Hengshui (HS), Baoding (BD), Tangshan (TS), Qinhuangdao (QHD), and Chengde (CD), representing four distinct ecological regions (Table 1 ). The latitudinal gradient ranged from 37°55′N (Jizhou, Hengshui) to 41°35′N (Weichang, Chengde). The distribution of the experimental locations is detailed in Table 1 . The waxy maize cultivar' Jingkenuo 768' (JKN768) was used as the test material. A randomized block design with three replications was used. A fertilizer application rate of 600 kg ha⁻¹was uniformly hand-applied to the corresponding plots before sowing and incorporated into the soil with a rotary tiller. The field was prepared by rotary tillage prior to sowing. Manual hill-drop sowing was conducted with a row spacing of 60 cm and a planting density of 52,500 plants ha⁻¹. All field management practices followed conventional methods used by local farmers. Table 1 The grain texture properties of waxy maize in different ecological regions Site Hardness (N) Cohesiveness (N/mm) Springiness (mm) Gumminess (N) Chewiness (mJ) Resilience HS 2.41c 0.06a 0.78a 0.24a 0.04b 0.05a BD 2.88b 0.0b3 0.84a 0.10c 0.11ab 0.01b TS 3.25a 0.03b 0.74a 0.12bc 0.11ab 0.01b QHD 2.66bc 0.05ab 0.85a 0.16b 0.16a 0.01b CD 2.61bc 0.04ab 0.70a 0.11bc 0.09ab 0.01b Data are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p < 0.05). Grain texture properties At 23 days after pollination, three ears were collected from each sampling site. The ears were steamed in an electric rice cooker, drained, and then packed in sealed bags to maintain a temperature of 60°C until testing. The measurement was performed using a texture analyzer (TMS-Pro, Japan), following the described method [ 19 ]. The instrument parameters were set as follows: a P/36 R probe, 20% compression ratio, compression speed of 1 mm/s, trigger force of 0.049 g, lift height of 60 mm, and test speed of 120 mm/min. Each sample was measured in triplicate. Soluble sugar and starch content At 23 days after pollination, three ears were collected from each sampling site. Kernels from the middle and lower sections of each ear were ground into fine powder. The soluble sugar content was determined using the anthrone-sulfuric acid colorimetric method [ 20 ]. The amylose and amylopectin contents were measured according to the instructions provided with the Solarbio assay kits (BC4260 and BC4270). Starch extraction and preparation Starch extraction and preparation At 23 days after pollination, three ears were collected from each sampling site, and 100 g of fresh kernels from the middle and lower sections of each ear were taken. Starch extraction and preparation were performed following the described method [ 1 ]. The fresh kernels were soaked in deionized water at 4°C for 48 h, then pulped using a juice extractor (model JYL-C23) and filtered. The filter residue was transferred to centrifuge tubes, and excess deionized water, absolute ethanol, and absolute diethyl ether were sequentially added. Each mixture was shaken for 10 min and then centrifuged at 4000 rpm for 10 min. This washing and centrifugation cycle was repeated three times. The resulting starch was dried at 40°C, passed through a 200-mesh sieve, and stored in a constant-temperature, constant-humidity chamber for subsequent analysis of its physicochemical properties. Scanning electron microscopy observation The sample stage was maintained with an appropriate quantity of starch granules. After gold coating using an ion sputter coater, the samples were observed with a scanning electron microscope (SU8010, HITACHI, Japan). Photographs were taken at magnifications of 1000× and 5000× [ 1 ]. Observation with optical and polarized light microscopy 10 mg sample was weighed and placed into 2 mL centrifuge tube, and 1 mL deionized water was added, and the mixture was vortexed to prepare 1% (w/v) starch suspension. A drop of this suspension was pipetted onto a glass slide pre-treated with a drop of 50% glycerol solution. Morphological analysis was performed using a polarized light microscope, with images captured under both bright-field and polarized light modes. Starch particle size distribution The volume distribution of starch particles was analyzed using a laser diffraction particle size analyzer (Mastersizer 3500, Malvern, England)[ 21 ]. This instrument measures sizes ranging from 0.1 to 2000 µm. The size distribution is expressed as the volume of equivalent spheres. The mean particle size is defined as the volume-weighted average. Each sample was measured in triplicate. Starch X-ray Diffraction Analysis XRD patterns of the starch samples were recorded using a D8 Advance X-ray diffractometer [ 22 ]. The instrument was operated at 200 mA and 40 kV. The diffraction angle (2θ) was scanned from 3° to 40° with a step size of 0.04° and a dwell time of 0.6 seconds per step. The relative crystallinity, defined as the ratio of the crystalline peak area to the total diffraction area, was calculated using MDI Jade 6.5 software. Each sample was measured in triplicate. Fourier transform infrared spectroscopy The ordered structure at the starch surface was determined using a Fourier transform infrared spectrometer (FTIR, Vertex 70, Bruker, Germany) according to the protocol described [ 22 ]. The starch sample was evenly distributed on the metal mold surface of an attenuated total reflectance accessory and then exposed to an infrared beam in the range of 4000 to 400 cm⁻¹. Spectra were collected with 32 scans at a resolution of 4 cm⁻¹. The spectral data within the 1200–800 cm⁻¹ range were analyzed using OMNIC 8.2 software. Each sample was measured in triplicate. Starch pasting properties The pasting properties of starch were determined using a Rapid Visco Analyzer (RVA4500, Perten, Australia). Precisely 1.96 g of starch was weighed and mixed with 26.04 g of ultrapure water to prepare a 28.00 g starch slurry at 7% (w/w). The testing profile was as follows: the sample and corresponding distilled water were first held at 50°C for 1 min, then heated from 50 to 95°C for 3.7 min, maintained at 95°C for 2.5 min, cooled to 50°C for 3.5 min, and finally held at 50°C for 2 min [ 1 ]. Data was analyzed using the Thermal Cycle for Windows software. Each sample was measured in triplicate. Starch thermal properties Thermodynamic properties of starch were determined using a differential scanning calorimeter (DSC Model 200 F3 Maia, NETZSCH, Germany). Starch was mixed with distilled water at a 1:2 (g:g) ratio, sealed in an aluminum pan, and kept overnight. Using an empty sealed aluminum crucible as a reference, the temperature was increased from 20 to 100°C at a rate of 10°C min⁻¹. The thermodynamic parameters were recorded and calculated [ 21 ]. After storing the samples at 4°C for 7 days, the retrogradation value and related parameters were measured under identical conditions. Each sample was analyzed in triplicate. Starch iodine binding capacity, blue value, and maximum absorption wavelength Approximately 80 mg of starch was weighed and added to 10 mL of a 50 mmol/L phosphate buffer (pH 7.0). The mixture was placed in a boiling water bath, shaken for 60 min, and then cooled to room temperature. 0.05 mL aliquot of the resulting solution was mixed with 4.85 mL of deionized water and 0.1 mL of iodine reagent. After rapid shaking, the mixture was allowed to stand for 15 min. Absorbance was measured using a UV-Vis spectrophotometer over a wavelength range of 500−700 nm. The iodine binding capacity was expressed as the ratio of absorbance at 635 nm to that at 520 nm (A635/A520) [ 23 ]. Each sample was analyzed in triplicate. Starch light transmittance The light transmittance of the starch paste were determined with slight modifications[ 24 ]. starch suspension (1%, w/v) was heated in a boiling water bath for 15 min with continuous stirring. After cooling at 25°C for 1 h, the suspension was stored at 4°C for 72 hours. The light transmittance was measured at 620 nm using an ultraviolet-visible spectrophotometer (UV-1700, Shimadzu, Japan) at 0, 24, 48, and 72 h, with distilled water serving as the blank. Each sample was measured in triplicate. Starch solubility and swelling degree The solubility and swelling degree were determined with slight modifications to the described method [ 25 ]. Precisely 160 mg starch sample (m0) was weighed into a pre-weighed 10 mL centrifuge tube (m1). Then, 8 mL of ultrapure water was added. The mixture was subjected to oscillatory heating in a water bath at 50, 60, 70, 80, and 90°C for 30 min, respectively. After cooling at room temperature, the sample was centrifuged at 3000×g for 10 min, and the precipitate weight (m2) was recorded. The supernatant was transferred to another pre-weighed 10 mL centrifuge tube (m3) and evaporated to dryness. The total mass (m4) was then measured. Each sample was analyzed in triplicate. The calculation formulas are as follows: Solubility (%) = (m4 / m0) × 100% (1) Swelling degree (g/g) = (m2 / m0) × 100% (2) Statistical analysis Mean values were calculated using Excel 2021. Analysis of variance and correlation tests between datasets were performed using the Statistical Package for the Social Sciences software (version 22.0). Differences between groups were determined for significance using the Least Significant Difference method. Figures were generated using Origin 2021. Correlation analyses were performed using the R software environment. Results Soil basic fertility The basic fertility of the 0−20 cm soil layer at in different sites are presented in Table 1 . The soil pH of HS and BD sites was 7.43, indicating a slightly alkaline condition. The pH values at TS, QHD, and CD sites ranged from 7.04 to 7.25. In terms of soil organic fertilizer, the organic matter content at the QHD site was significantly higher than at other sites, reaching 19.27 mg/kg. In contrast, the TS site had the lowest soil organic matter content of 10.09 mg/kg. The available soil nitrogen content followed the order: QHD > HS > BD > CD > TS. The QHD site exhibited a soil-available nitrogen content of 84.23 mg/kg, whereas the TS site had 54.83 mg/kg, indicating a significantly weaker nitrogen supply capacity than in other regions. Analysis of soil available phosphorus and potassium contents across different sites revealed that the HS site had the highest levels of both nutrients, with identical values of 604.67 mg/kg. Furthermore, BD, TS, QHD, and CD sites exhibited a consistent trend in soil-available phosphorus and potassium. Specifically, the BD site recorded 101.00 mg/kg; TS and QHD sites measured 137.00 mg/kg and 133.00 mg/kg, respectively, the CD site had an available nutrient level identical to that of the QHD site. In summary, the sites exhibited significant variation in soil fertility. The QHD site exhibited sufficient soil organic matter and alkali-hydrolyzable nitrogen, the HS site demonstrated significantly higher available nutrient level. In contrast, the overall soil fertility at the TS site was relatively weak. This regional heterogeneity in soil fertility provide a valuable reference for regional cultivation nutrient management of waxy maize. Climatic ecological factors The climatic ecological factors for the period from May to August across different sites are presented in Table 2 . The highest temperatures at the HS site from May to August were significantly higher than those at other sites, reaching 40.33°C in June. In contrast, the CD site exhibited significantly low temperatures, with the lowest in May at 0.54°C and in August at 2.77°C, its overall temperature was lower than that of the other sites. The average temperature from May to August followed the trend of HS > TS > BD > QHD > CD. At the HS site, the monthly average temperature remained above 22°C, whereas at the CD site, the average temperature for July and August remained below 21°C. Analysis of the effective accumulated temperature indicated that the total effective accumulated temperature from May to August at the HS site reached 2117.28°C, the highest among all sites, with monthly accumulations exceeding 500°C (reaching 615.02°C in July). However, the total effective accumulated temperature at the CD site was 925.07°C, which was less than half that at the HS site. The effective accumulated temperature in May was approximately 66.35°C, the thermal conditions at this site were significantly weaker than those in other regions. Analysis of precipitation across the sites exhibited an uneven spatial distribution. TS and QHD sites experienced concentrated rainfall in August (252.96 and 250.05 mm, respectively). At the BD site, precipitation in July and August accounted for 75% of the total. The highest rainfall values recorded at HS and CD sites were 126.74 and 248.85 mm, respectively, in July. In summary, meteorological conditions varied significantly among the five sites. The HS site had sufficient heat; however, relatively low rainfall. In contrast, the CD site was characterized by insufficient heat and a more uniform precipitation distribution. TS and QHD sites exhibited concentrated rainfall in the later growth period. This regional heterogeneity in ecological factors provides fundamental data to support research on the regional adaptability of waxy maize. Table 2 The grain nutritional quality of waxy maize in different ecological regions Site Sugar (mg/g) Total starch (mg/g) Amylopectin (mg/g) Amylose (mg/g) Amylopectin proportion(%) HS 79.58b 429.49c 428.60b 0.89c 99.79a BD 87.14a 456.57a 455.65b 0.92a 99.80a TS 84.21ab 458.10a 457.27b 0.83a 99.82a QHD 82.01ab 442.81b 441.41a 1.41b 99.68a CD 73.31c 455.32a 454.37b 0.96a 99.79a Data are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p < 0.05). Grain texture properties Across the different sites, significant differences (P < 0.05) in the texture characteristics of waxy maize kernels were observed (Table 3 ). Grain hardness was highest at the TS site, followed by the BD site, and lowest at the HS site. Grain cohesiveness was highest at the HS site, while BD and TS sites exhibited lower values. The grain elasticity exhibited no significant difference (P > 0.05) across sites, suggesting that the experimental location did not affect it. Grain adhesiveness was significantly higher (P < 0.05) at the HS site than at BD and TS sites, with no significant difference between BD and TS sites. Grain chewiness was highest at the QHD site and lowest at the HS site; the remaining sites exhibited a medium level with no significant differences among them. Table 3 The volume distribution and average granule size of waxy maize starch in different ecological regions Site D(4,3) D(3,2) Volume distribution (µm) 15µm HS 12.79e 8.33e 9.39a 46.48a 44.14e BD 14.45d 9.37c 8.00c 34.69c 57.31d TS 22.49a 11.08a 6.62e 26.06e 67.32a QHD 15.00b 9.27d 8.57b 33.75b 57.69c CD 14.81c 10.00b 7.06d 32.05d 60.88b Data are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p < 0.05). Grain nutritional quality The grain nutritional quality of waxy maize exhibited significant differences across the sites (Table 4 ). The soluble sugar content in the grains was significantly higher at the BD site compared to HS and CD sites (P < 0.05). TS and QHD sites exhibited intermediate values, indicating no significant difference from the BD site, while the CD site recorded the lowest sugar content. Regarding total starch content, the TS site had the highest level, the BD and CD sites did not differ significantly from TS, Thses three sites had significantly higher total starch content than QHD and HS sites (P < 0.05), with the HS site exhibiting the lowest value. For amylose content, the QHD site had the highest level, which was significantly greater than that at all other sites (P < 0.05). The HS site exhibited the lowest amylose content, and no significant differences were observed among BD, TS, and CD sites. The amylopectin content in the grains was significantly higher at the QHD site compared to the other sites (P > 0.05). Table 4 The pasting properties of waxy maize starch in different ecological regions Site PV(cP) TV(cP) FV(cP) BD(cP) SB(cP) Ptemp(°C) HS 1441.3b 664.7c 772.0d 760.0c 107.3a 77.8a BD 1859.3a 871.0a 941.7b 988.3a 70.7a 68.7a TS 1824.7a 906.0a 976.3a 918.7ab 70.3a 77.5a QHD 1340.3b 533.0d 637.3e 807.3bc 104.3a 76.7a CD 1788.0a 795.7b 876.7c 992.3a 81.0a 74.0a Data are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p < 0.05). Starch granule morphology, optical microscopy, and polarized cross structure The scanning electron microscope (SEM) plays a crucial role in characterizing the structure of starch granules. SEM images of starch from different sites revealed potential variations in the size and shape of starch granules across treatments. Starch granule size significantly influences the quality characteristics of isolated starch. The starch granules from the HS site were relatively small and uniformly distributed, with relatively smooth surfaces (Figs. 1 -A-B). In contrast, the starch granules from the TS site were significantly larger than those from other sites, with nearly spherical shapes and plump surfaces. This observation aligns with the results described in Table 5 , which indicates that the TS site has the largest volume average particle size. Optical microscopy (Fig. 1 -C) indicated that the waxy corn starch granules from the HS site were well dispersed with minimal aggregation. Conversely, the starch granules from the TS site exhibited a higher degree of aggregation, attributed to stronger surface forces associated with larger particle sizes. In polarized-light micrographs (Fig. 1 -D), waxy corn starch from all sites exhibited the typical Maltese cross extinction pattern, indicating the presence of crystalline starch. The extinction cross of starch from the TS site exhibited greater clarity, consistent with its highest crystallinity characteristic as presented in Table 3 . In contrast, the extinction cross of starch from the QHD site appeared relatively indistinct, consistent with its moderate starch crystallinity. Table 5 The thermal properties of waxy maize starch in different ecological regions Site To(℃) Tp(℃) Tc(℃) ΔH(J/g) R(%) HS 70.52b 78.53ab 84.67a 4.03d 32.22b BD 70.75b 79.33ab 85.17a 6.31b 24.94c TS 81.30a 83.20a 88.76a 7.22a 30.21b QHD 70.69b 73.63b 76.62b 4.62c 38.79a CD 65.32b 77.56ab 79.84b 4.06d 22.29c Data are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p < 0.05). Starch size distribution There were significant differences in the average particle size and volume distribution of waxy corn starch among different sites (Table 5 ). Regarding the average starch granule size, both the volume-average particle size D (4,3) and the surface-area mean diameter D (3,2) at the TS site were significantly higher than those at other sites ( 15 µm) reached 67.32%, significantly higher than at other sites, while the proportion of medium starch particles (5–15 µm) was only 26.06%. These results indicate that large particles dominated the waxy maize starch granules at the TS site, whereas those at the HS experimental site were primarily small particles. The starch particle size volume distribution curve (Fig. 2 -A) indicates that the peak of the starch size distribution at the TS site shifted towards the large particle size range. In contrast, the distribution peak at the HS experimental site was concentrated in the small-particle size range. This observation aligns with the results in Table 5 , where the volume-average particle size was largest at the TS experimental site and smallest at the HS site, providing a visual demonstration of the differences in particle size among the sites. Starch relative crystallinity and short-range ordered structure The crystallinity of starch is associated with the ordered structure of amylopectin molecules within the granules. XRD patterns (Fig. 2 -B) revealed that starch from all sites exhibited typical type A crystalline characteristics, with diffraction peaks at 15°, 17°, 18°, and 23°. Differences in starch relative crystallinity were observed among the sites. The TS site exhibited significantly higher relative crystallinity than the other sites (P < 0.05), the HS site had the lowest value. The remaining sites exhibited no significant differences.In the FTIR full spectra (Fig. 2 -C), the absorption peak trends in regions, including 3400 cm⁻¹ (hydroxyl stretching vibration), 2930 cm⁻¹ (C-H stretching vibration), and 1640 cm⁻¹ (water molecule bending vibration), were consistent across all sites. However, differences in IR ratios were observed within the characteristic peak region (Fig. 2 -D). The 1045/1022 ratio at the HS site was significantly lower than that at the other sites (P < 0.05), the 1022/995 ratio exhibited no significant differences among the sites. Starch pasting properties The peak viscosity (PV) of starch from the BD and TS sites was significantly higher than that from HS and QHD sites presented in Table 6 . The trend of trough viscosity (TV) aligned with that of final viscosity. The QHD site exhibited the lowest TV. The breakdown value reflects the thermal stability of starch. The breakdown values for BD and CD sites were significantly higher than those for the HS site (P < 0.05). Furthermore, no significant differences were observed in setback value and starch pasting temperature among the sites. Table 6 The iodine binding capacity, blue value and maximum absorption wavelength of waxy maize starch in different ecological regions. Site Iodine binding capacity Blue value Maximum absorption wavelength(nm) HS 0.031b 0.887b 547.00ab BD 0.072b 1.727a 553.75a TS 0.072b 1.898a 542.25bc QHD 0.159a 1.897a 537.00c CD 0.079b 2.359a 540.25bc Data are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p < 0.05) Starch thermal properties The QHD site exhibited a smaller area under the starch heat flow peak(Table 7), corresponding to its lower pasting enthalpy change. This visually demonstrates the differential regulation of starch thermal pasting properties across the sites. Specifically, the ΔHgel was highest at the TS site, significantly exceeding that at HS, QHD, and CD sites (P < 0.05). HS and CD sites recorded the lowest ΔHgel values. The retrogradation value was significantly higher at the QHD site than at the other sites (P < 0.05), whereas BD and CD sites exhibited lower values. This suggests that the starch at the QHD site is more susceptible to pasting. Starch iodine binding capacity, blue balue, and maximum absorption wavelength The starch from the QHD site exhibited significantly higher (P < 0.05) iodine binding capacity than that from the other sites (Table 8). HS, BD, TS, and CD sites indicated no significant differences. This indicates that the starch from QHD possesses a stronger ability to bind iodine, which can be associated with its amylose content or molecular chain structure. This finding aligns with the result presented in Table 4 , which indicates that the QHD site had the highest amylose content. The blue value of starch from the HS site was significantly lower than that from the other sites (P < 0.05). BD and CD site both exhibited relatively high blue values with no significant difference between them. The blue value further reflects differences in the color development intensity of the starch-iodine complex, and its variation trend is consistent with that of the iodine binding capacity. The BD site exhibited the highest maximum absorption wavelength for starch, while the QHD site demonstrated the lowest, with the QHD site being significantly lower than the BD site (P < 0.05). Additionally, no significant differences were observed among the remaining sites. Starch light transmittance The starch light transmittance at each site decreased with prolonged storage time at 4°C (Fig. 3 -A). The transmittance at the TS site was significantly higher than that at the other sites at all time sites (P < 0.05). At the TS site, the starch light transmittance reached 80% at 0 and 24 h; however, it was reduced below 80% at 48 and 72 h. At the other sites, the initial transmittance values at various time sites ranged from 60% to 75%. At 0 h, the starch light transmittance at the TS site was significantly greater than that at the other sites, while no significant differences were observed among the other sites. At 72 h, the TS site remained significantly higher than HS and QHD sites; however, it was not significantly different from BD and QHD sites. The TS site exhibited the largest decline in starch light transmittance, suggesting that its starch paste had better initial transparency but weaker long-term stability. Starch solubility and swelling power The starch solubility at all sites increased with increasing temperature (Fig. 3 -B). Solubility and swelling power exhibited a significant correlation with temperature, a phenomenon attributed to the increased mobility of starch and water molecules, with higher temperatures promoting the diffusion of amylose and amylopectin. When the temperature reached 90°C, the starch swelling power at the TS site was significantly higher than that at the other sites (P < 0.05). This finding is associated with the starch structural characteristics of the TS site (large particle size and high crystallinity) as presented in Table 4 . On the one hand, the larger particle size provides more physical space for the starch, resulting in a greater expansion volume after water absorption. On the other hand, the hydrogen bond network in the crystalline region of starch with high crystallinity undergoes extensive dissociation at high temperatures. The strong water absorption driving force during this dissociation process allows it to bind more water. In contrast, the starches from other sites (HS and BD) have smaller mean particle sizes and lower relative crystallinity, lacking sufficient swelling space and exhibiting a weaker water-absorption driving force. Consequently, their starch swelling power was significantly lower than that of the TS site. The solubility of starch at each site increased with increasing temperature (Fig. 3 -C). Within the 50–90°C temperature range, starch solubility at all sites increased progressively with increasing temperature. A greater temperature rise corresponded to a more pronounced rate of increase in solubility. When the temperature reached 90°C, starch solubility at each site peaked; however, the differences were not statistically significant. Correlation analysis between grain texture and starch physicochemical properties Pearson correlation coefficients and mantel test results for the relationships between grain texture and starch physicochemical properties are presented in Fig. 4-A. Starch granule size positively regulates grain texture. Stickiness and chewiness exhibited strong positive correlations with small starch granules, indicating that the tested parameters have a relatively lesser influence on cohesiveness. The associations between indicators, including Tp and Tc, and all texture characteristics were not statistically significantly, indicating that these parameters were not key factors in regulating grain texture in this study. In summary, starch granule size is a core indicator regulating grain stickiness, chewiness, and hardness, R% primarily influences grain elasticity. These results provide a quantitative basis for targeted improvement in grain texture quality by optimizing starch physicochemical properties. Principal component analysis of soil conditions, meteorological factors, and grain quality The results of principal component analysis (Fig. 4-B) indicated that the first two principal components (PC) cumulatively explained 58.8% of the total variation in grain quality-associated indicators of waxy maize across different ecological regions in northern China. PC1, accounting for 34.9% of the variance, represented the dimension of structural strength and thermal stability, its positive direction was significantly associated with variables, including the proportion of large-sized starch (> 15 µm), relative crystallinity of starch, peak gelatinization viscosity, grain hardness, and soil available nitrogen content. Conversely, the negative direction was associated with variables including the proportions of small sized starch (< 5 µm and 5–15 µm), adhesiveness, chewiness, rainfall, and retrogradation value. PC2, explaining 23.9% of the variance, corresponded to the dimension of starch composition and pasting properties, its positive direction was associated with high amylose content, a strong setback tendency, and a relatively high pasting temperature. Discussion This study indicates that differences in grain texture of waxy maize across various ecological regions are primarily due to the regulatory effects of environmental factors on starch granule morphology and crystal structure. Specifically, the TS site exhibited the highest kernel hardness, whereas the HS site demonstrated the lowest hardness and relatively high adhesiveness. This phenomenon is closely associated with the starch granule size distribution, the TS site was dominated by large-granule starch (> 15 µm), while the HS site was rich in small-granule starch (< 5 µm). Multivariate correlation analysis (Fig. 6) further revealed that the proportion of small granule starch (< 5 µm) exhibited a significant positive correlation (P ≤ 0.01) with both adhesiveness and chewiness of the kernels. The underlying mechanism is associated with the greater specific surface area of small granule starch, which promotes the interactions between granules and molecules, thereby contributing to a more pronounced sticky taste during mastication. This observation aligns with the conclusion that increasing the proportion of small granule starch improves the viscoelasticity of food products [ 26 ]. Conversely, large granule starch can provide a more supportive physical structure [ 27 ]. The TS site not only exhibited a larger starch granule size but also the highest relative crystallinity. The synergistic effect of these two factors formed a dense crystalline network, significantly increasing the mechanical hardness of the kernels. This finding is consistent with the established theory that both starch crystallinity and granule size collectively determine hardness [ 28 ]. However, the HS site was characterized by a predominance of small granule starch and low crystallinity, resulting in loose granule binding and, consequently, reduced hardness [ 29 ]. The crystalline structure of starch is a critical factor that interacts with environmental factors to determine its final functional properties. XRD and FTIR analyses consistently indicate that starch from the TS site exhibits the highest crystallinity, while that from the HS site demonstrates the lowest. This variation correlates positively with particle size, as larger granules provide more space for the orderly arrangement of starch molecular chains, which facilitates the formation and maintenance of well-defined crystalline regions [ 30 ]. In contrast, the crystalline structure of small granule starch tends to be looser and less complete [ 31 ]. The lower 1045/1022 cm⁻¹ FTIR peak intensity ratio observed for the HS site further confirms its insufficient crystalline order. These differences in crystallinity directly determine the thermodynamic behavior of the starch. ΔH represents the energy required to disrupt the double helix structure within the starch crystalline region [ 32 ]. The high crystallinity of starch from the TS site suggests a denser, more stable internal hydrogen-bond network, thereby requiring greater energy input for gelatinization, which aligns with findings reported in rice studies [ 33 ]. The higher PV values observed at TS and BD sites are due to the starches at these sites, which possess both large granules and high crystallinity. Large granule starch exhibits a stronger water absorption and swelling potential during the initial stage of gelatinization, while a highly crystalline structure offers greater resistance to heat shear. The synergistic effect of these two factors [ 34 ] allows starch to accumulate and reach a higher viscosity peak during heating [ 35 – 36 ]. Environmental conditions modify the chemical composition and molecular structure of starch by regulating the starch biosynthesis pathway, thereby affecting its functional properties. In this study, the TS site exhibited the highest total starch content, likely because its favorable environment effectively promoted starch synthase activity and substance accumulation [ 21 , 37 ]. Conversely, waxy maize at the HS site exhibited the lowest total starch content, which can result from suppression of starch synthesis gene expression under environmental stress. Notably, the total starch content of waxy maize at the QHD site was not the highest; however, its amylose content was significantly higher than at other sites, resulting in a substantial increase in setback viscosity. This is attributed to the greater tendency of amylose molecules to rearrange and form a hard gel during the cooling process [ 38 ]. Furthermore, the strongest iodine-binding capacity of waxy maize starch from the QHD site directly confirmed its higher amylose content [ 39 ]. The lower starch blue value at the HS site reflected its lower amylose content. Differences in the λmax of starch across the sites suggested that environmental conditions could influence fine molecular structures, including the amylopectin branch-chain length. The starch granule and crystal structure indirectly determine its performance after gelatinization. The highest initial transmittance of waxy maize starch from the TS site could be attributed to a more uniform gel network formed after gelatinization of its large, highly crystalline granules. Waxy maize at the TS site exhibited the highest starch swelling power at 90°C because the high temperature sufficiently disrupted its dense crystalline structure, resulting in significant dissociation of hydrogen bonds, while the large granules provided additional space to bind more water [ 35 ]. Furthermore, the higher kernel soluble sugar content observed at the BD site can be attributed to lower efficiency in converting photosynthetic products into starch under its specific environmental conditions, including temperature and moisture, resulting in sugar accumulation. This indicates environmental plasticity in the distribution of carbon metabolic fluxes. Meteorological factors exhibited significant spatial heterogeneity in shaping the quality of waxy maize kernels. In regions represented by the HS site, the grain-filling stage (May to August) was characterized by abundant heat resources, with an effective accumulated temperature of 2117.28°C; however, relatively low precipitation of 333.90 mm. The favorable thermal conditions likely accelerated starch biosynthesis, promoted the formation of small starch granules, and optimized grain viscoelasticity. In contrast, the CD site experienced a lower effective accumulated temperature of 925.07°C during grain filling, with evenly distributed rainfall. However, the low temperatures likely inhibited starch synthase activity, resulting in insufficient starch granule development and, consequently, reduced grain adhesiveness and chewiness. These findings align with existing research, indicating that high-temperature treatment induces irregular surface morphology and increases the average starch granule size of waxy maize starch, while reducing its gelatinization viscosity and enthalpy value [ 40 ]. Conversely, moderate low temperature decreases the average starch granule size [ 26 ], increases the proportion of short chains in amylopectin, and reduces relative crystallinity, ultimately impacting grain stickiness. Similar studies in rice have reported that sustained high temperatures modify starch component accumulation, particle morphology, and crystal structure, negatively impacting edible quality [ 41 ]. Conversely, low-temperature stress increases amylose content and retrogradation value and modifies the fine structure of amylopectin [ 42 ]. Furthermore, TS and QHD sites experienced heavy rainfall (> 250 mm) during the late grain-filling stage (August), which induced grain metabolic water stress. This disturbance could have disrupted the normal construction and stability of starch structure, thereby affecting the final texture. This observation is consistent with previous reports that waterlogging stress reduces starch content and modifies its gelatinization properties [ 43 ]. Soil conditions are another key driver of regional differentiation in waxy maize quality. In this study, the soil at the QHD site was rich in organic matter and readily available nitrogen. Therefore, the grains exhibited significantly higher amylose and amylopectin contents, as well as a higher iodine-binding capacity. However, the clarity of the starch polarization cross was poorer, and gelatinization stability was weaker, sufficient nitrogen supply can promote starch synthesis, and thus affect starch properties to a certain extent [ 44 ]. The soil at the HS site had higher levels of readily available phosphorus and potassium, indicating a significant nutrient supply potential. However, the grains exhibited the lowest total starch content, relative crystallinity, and swelling power. The starch granules were predominantly small, smooth, and well dispersed. This suggests that elevated soil phosphorus and potassium levels can have inhibited the full development and orderly aggregation of starch granules [ 45 ]. At the TS site, the soil had low organic matter content and poor nitrogen supply capacity.The starch exhibited the largest average granule size, the highest crystallinity and enthalpy value, as well as excellent thermal stability of gelatinization. Additionally, the grain hardness was relatively higher. This reveals an interesting phenomenon, low nitrogen soil conditions promote the development of waxy maize starch granules toward larger size and higher crystallinity, forming a more compact starch granule structure [ 46 ]. Soil fertility at the BD site was medium. The grains exhibited higher kernel soluble sugar content, PV, and breakdown value; however, the starch's thermal stability was only moderate. Previous studies have suggested that sandy soil has a poor water-retention capacity under drought conditions, thereby exacerbating the inhibitory effect of water stress on starch synthesis. In contrast, the stronger water-retention capacity of clay soil can buffer the adverse effects of excessive water on the stability of the starch structure under rainy conditions [ 43 ]. This provides a theoretical basis for explaining the quality differences among different soil types within the same climatic region. Conclusion The regional heterogeneity of meteorological, ecological, and soil factors constitutes the core determinant of waxy maize grain quality and starch properties. The hydrothermal combination characteristics in different ecological regions exert significantly distinct regulatory effects on starch granule development, crystalline structure, and functional properties. At the HS site in the central-southern Hebei region, abundant heat supply coupled with precipitation scarcity during the growth period shaped starch characteristics, resulting in small particle size and low crystallinity, conferring a quality advantage in higher grain adhesiveness. In BD, concentrated rainfall during the later growth stage resulted in starch granules of medium size and medium crystallinity. This resulted in higher grain hardness and soluble sugar content, albeit with weaker starch thermal stability. In the northern Hebei region, insufficient heat supply at the CD site inhibited the orderly development of starch granules, directly resulting in overall quality deterioration in waxy maize. The regulatory effects of concentrated precipitation during the later growth stage differed significantly between the two experiment sites in the Eastern Hebei region. At the TS site, a starch crystal structure characterized by large particle size and high crystallinity formed, resulting in synergistic increases in grain hardness, total starch accumulation, and starch PV. In contrast, the QHD site exhibited a higher proportion of amylose in the grains and improved iodine-binding capacity, thereby strengthening the starch retrogradation characteristics. Therefore, the waxy maize from this site exhibited unique aging resistance and gel functional properties. This study clarifies the key influencing factors and regulatory mechanisms underlying quality formation in waxy maize across different ecological regions of Hebei Province. This study provide precise theoretical support for the regional cultivation of high-quality waxy maize and the breeding of special varieties. Declarations Funding This study was financially supported by the National Key Research and Development Program of China (Grant No.2023YFD2301505). 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Zhang","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA5ElEQVRIiWNgGAWjYDACCQjFw8/ekPggocKGeC0ykj0HHhs8OJNGvBYbgxuOzyQfth0irEN+dvOzh1/b7HgMbjCnVSSwHWDgb+9OwKuFcc4xc2PZtmQeydttaTcSeO4wSJw5uwGvFmaJBDNpybYDPHx3zgC1SDxjMJDIxa+FTSL9G1gLw438bwUJBocJa+GRyDGT/AjUInAjIY0hIYEILRISOWXSDOeAfuk5kCyRcCCNh6Bf5Gekb5P8UWZnD4rKjz//2cjxt/fi1wICzLxsSC4lqBwEGH/8IUrdKBgFo2AUjFQAAFyPS3CCBQEqAAAAAElFTkSuQmCC","orcid":"","institution":"Hebei Agricultural University","correspondingAuthor":true,"prefix":"","firstName":"Yuechen","middleName":"","lastName":"Zhang","suffix":""}],"badges":[],"createdAt":"2026-03-21 04:38:17","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-9183115/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-9183115/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":106240201,"identity":"abd5af83-4689-4239-a4c3-f0e48ae0c9e8","added_by":"auto","created_at":"2026-04-06 14:45:09","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":276908,"visible":true,"origin":"","legend":"\u003cp\u003eThe scanning electron microscopy(A-B), optical microstructure(C) and polarized light cross structure (D) of waxy maize starch in different ecological regions\u003c/p\u003e","description":"","filename":"1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9183115/v1/ff153668cbade369683a3638.jpg"},{"id":106404124,"identity":"27229041-180c-4283-9153-d5639aa236b3","added_by":"auto","created_at":"2026-04-08 09:15:31","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":167453,"visible":true,"origin":"","legend":"\u003cp\u003eThe granule size distribution (A), crystalline properties (B), infrared spectrum (C) and characteristic peak ratio (D) of waxy maize starch in different ecological regions\u003c/p\u003e","description":"","filename":"2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-9183115/v1/fa140b8af612f2513b9631d4.jpg"},{"id":106240203,"identity":"5388b640-125d-44cf-a30a-fbd2bd90faef","added_by":"auto","created_at":"2026-04-06 14:45:09","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":105215,"visible":true,"origin":"","legend":"\u003cp\u003eThe light transmittance (A), swelling power (B) and solubility (C) of waxy maize starch in different ecological regions\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-9183115/v1/7c4d1aa1bdd75794d8f8bd60.png"},{"id":106406018,"identity":"59475d6b-25f7-4777-a013-d33b3ff1e2c1","added_by":"auto","created_at":"2026-04-08 09:29:26","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1951559,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-9183115/v1/5b779d8c-eddb-42d5-af8f-16655ab5a3e6.pdf"},{"id":106240200,"identity":"c02d9f13-5f2f-43bc-9ec8-cef0041d1db7","added_by":"auto","created_at":"2026-04-06 14:45:09","extension":"doc","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":821956,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryMaterial.doc","url":"https://assets-eu.researchsquare.com/files/rs-9183115/v1/6d920543f34442eb619ac680.doc"}],"financialInterests":"No competing interests reported.","formattedTitle":"Variations in grain textural and starch physicochemical properties of waxy maize across contrasting ecological regions","fulltext":[{"header":"Introduction","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eWaxy maize is a distinctive type of maize extensively cultivated in China, its starch is almost entirely composed of amylopectin, which provides high viscosity, a low retrogradation rate, and good stability [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. These unique physicochemical characteristics of the starch give waxy maize a high value for processing and utilization, resulting in its extensive application in the food industry and in industrial products, including thickeners, emulsifiers, and adhesives [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. As the primary component of waxy maize, starch is a key determinant of its quality. Specifically, the physicochemical properties of the starch are directly influenced by the ratio and structure of amylose to amylopectin, which, in turn, determine its suitability for specific applications [\u003cspan additionalcitationids=\"CR4\" citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eStarch possesses a multi-scale hierarchical structure, characterized by features, including amylopectin chain length, particle morphology, crystalline structure, and molecular conformation. Key indicators for evaluating waxy corn starch properties include amylopectin content, particle size distribution, crystallinity, and gelatinization temperature. Notably, these characteristics are susceptible to environmental and cultivation factors during the grain formation period, including temperature, waterlogging stress at the flowering stage, and nitrogen level [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e]. Furthermore, exogenous substances, including coronatine and salicylic acid, can improve waxy corn grain yield and starch quality by regulating the activity of key enzymes involved in starch synthesis and the expression of related genes [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe growth environment is a critical regulatory factor in starch macromolecule biosynthesis. Studies indicate that temperature fluctuations during the grain-filling stage can affect the activities of starch synthase and branching enzymes, thereby altering the amylopectin chain-length distribution and crystal packing modes [\u003cspan additionalcitationids=\"CR10\" citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. Drought stress reduces the starch content in waxy maize, increases the levels of total protein, globulin, and glutelin, and results in an increase in starch average particle size, amylopectin average chain length, and relative crystallinity. Furthermore, the impact of drought stress is more pronounced during the kernel-formation stage than during the grain-filling and enrichment stages [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e]. Variations in the ecological region's environmental conditions contribute to changes in starch properties. A study comparing broomcorn millet starch from Yangling and Yulin in Shaanxi Province observed that in the warmer Yangling region, the starch exhibited a higher proportion of long amylopectin chains and greater relative crystallinity. This was accompanied by an increase in gelatinization temperature, increased shear resistance, and a decrease in retrogradation value [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. Furthermore, a study on waxy maize has confirmed that high-temperature stress during kernel formation increases the average particle size of starch granules and the proportion of long amylopectin chains, thereby altering their gelatinization and thermodynamic properties [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e]. A comparative study of heat-sensitive and heat-tolerant waxy corn varieties revealed that high temperatures during grain development result in surface depressions on starch granules, increased particle size, a higher proportion of long amylopectin chains, and elevated relative crystallinity. Subsequently, these changes reduce the gelatinization viscosity and enthalpy of starch while increasing the retrogradation rate [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eHistorically, waxy maize was considered a specialty crop. However, due to its unique starch composition, strong processing adaptability, and outstanding nutritional value, it has now emerged as a potential crop for diversifying food industry raw materials and increasing the added value of agricultural products. Waxy maize is extensively cultivated across China, spanning multiple ecological regions from the humid, hot south to the arid north. These regions exhibit significant differences in temperature, precipitation, and soil fertility. Variations in these natural ecological conditions can result in differences in quality within the same variety grown in different regions. Additionally, the extent of these differences can be comparable to that observed among different varieties within the same area [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Therefore, this study hypothesizes that waxy maize produced across different ecological regions can exhibit significant variation in grain texture characteristics and in the physicochemical properties of starch. These differences would directly affect the potential of waxy maize in both food and non-food industries, as the development of specific products depends on starch raw materials with the corresponding properties.\u003c/p\u003e \u003cp\u003eThis study used the novel fresh-eating waxy maize variety \"Sweet Waxy\" and its representative cultivar Jingkenuo 768 [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e] as experimental materials. These materials were cultivated across four ecological regions in Hebei Province, characterized by contrasting ecological factors. This study aimed to identify patterns of grain quality variation in waxy maize across these regions by analyzing the relationships among grain texture characteristics, starch physicochemical properties, and environmental conditions in different ecological regions of Hebei Province. The findings can provide theoretical and technical references for high-quality waxy maize cultivation in the province. The specific objectives of this study were as follows: (1) analyzing the grain texture, starch particle morphology, crystalline and infrared spectral structural features of waxy maize from different ecological regions in Hebei Province and evaluating the functional properties of its starch; (2) investigating the association mechanisms between the grain texture of waxy maize and the structural and functional properties of its starch; (3) identifying the influences of meteorological ecological factors and soil conditions on the grain texture and physicochemical properties of starch in waxy maize.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eExperimental site conditions\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003eFive experimental sites were selected within Hebei Province, including Hengshui (HS), Baoding (BD), Tangshan (TS), Qinhuangdao (QHD), and Chengde (CD), representing four distinct ecological regions (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The latitudinal gradient ranged from 37\u0026deg;55\u0026prime;N (Jizhou, Hengshui) to 41\u0026deg;35\u0026prime;N (Weichang, Chengde). The distribution of the experimental locations is detailed in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. The waxy maize cultivar' Jingkenuo 768' (JKN768) was used as the test material. A randomized block design with three replications was used. A fertilizer application rate of 600 kg ha⁻\u0026sup1;was uniformly hand-applied to the corresponding plots before sowing and incorporated into the soil with a rotary tiller. The field was prepared by rotary tillage prior to sowing. Manual hill-drop sowing was conducted with a row spacing of 60 cm and a planting density of 52,500 plants ha⁻\u0026sup1;. All field management practices followed conventional methods used by local farmers.\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe grain texture properties of waxy maize in different ecological regions\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eHardness\u003c/p\u003e \u003cp\u003e(N)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eCohesiveness\u003c/p\u003e \u003cp\u003e(N/mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eSpringiness\u003c/p\u003e \u003cp\u003e(mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eGumminess\u003c/p\u003e \u003cp\u003e(N)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eChewiness\u003c/p\u003e \u003cp\u003e(mJ)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eResilience\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.41c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.06a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.78a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.24a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.04b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.05a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.88b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.0b3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.84a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.10c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.11ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.25a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.03b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.74a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.12bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.11ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eQHD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.66bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.05ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.85a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.16b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.16a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e2.61bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.04ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.70a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.11bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.09ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003eData are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eGrain texture properties\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eAt 23 days after pollination, three ears were collected from each sampling site. The ears were steamed in an electric rice cooker, drained, and then packed in sealed bags to maintain a temperature of 60\u0026deg;C until testing. The measurement was performed using a texture analyzer (TMS-Pro, Japan), following the described method [\u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. The instrument parameters were set as follows: a P/36 R probe, 20% compression ratio, compression speed of 1 mm/s, trigger force of 0.049 g, lift height of 60 mm, and test speed of 120 mm/min. Each sample was measured in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eSoluble sugar and starch content\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eAt 23 days after pollination, three ears were collected from each sampling site. Kernels from the middle and lower sections of each ear were ground into fine powder. The soluble sugar content was determined using the anthrone-sulfuric acid colorimetric method [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e]. The amylose and amylopectin contents were measured according to the instructions provided with the Solarbio assay kits (BC4260 and BC4270).\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e\n\u003ch3\u003eStarch extraction and preparation\u003c/h3\u003e\n\u003cdiv class=\"Heading\"\u003eStarch extraction and preparation\u003c/div\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eAt 23 days after pollination, three ears were collected from each sampling site, and 100 g of fresh kernels from the middle and lower sections of each ear were taken. Starch extraction and preparation were performed following the described method [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. The fresh kernels were soaked in deionized water at 4\u0026deg;C for 48 h, then pulped using a juice extractor (model JYL-C23) and filtered. The filter residue was transferred to centrifuge tubes, and excess deionized water, absolute ethanol, and absolute diethyl ether were sequentially added. Each mixture was shaken for 10 min and then centrifuged at 4000 rpm for 10 min. This washing and centrifugation cycle was repeated three times. The resulting starch was dried at 40\u0026deg;C, passed through a 200-mesh sieve, and stored in a constant-temperature, constant-humidity chamber for subsequent analysis of its physicochemical properties.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e\n\u003ch3\u003eScanning electron microscopy observation\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe sample stage was maintained with an appropriate quantity of starch granules. After gold coating using an ion sputter coater, the samples were observed with a scanning electron microscope (SU8010, HITACHI, Japan). Photographs were taken at magnifications of 1000\u0026times; and 5000\u0026times; [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e\n\u003ch3\u003eObservation with optical and polarized light microscopy\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003e10 mg sample was weighed and placed into 2 mL centrifuge tube, and 1 mL deionized water was added, and the mixture was vortexed to prepare 1% (w/v) starch suspension. A drop of this suspension was pipetted onto a glass slide pre-treated with a drop of 50% glycerol solution. Morphological analysis was performed using a polarized light microscope, with images captured under both bright-field and polarized light modes.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eStarch particle size distribution\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe volume distribution of starch particles was analyzed using a laser diffraction particle size analyzer (Mastersizer 3500, Malvern, England)[\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. This instrument measures sizes ranging from 0.1 to 2000 \u0026micro;m. The size distribution is expressed as the volume of equivalent spheres. The mean particle size is defined as the volume-weighted average. Each sample was measured in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eStarch X-ray Diffraction Analysis\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eXRD patterns of the starch samples were recorded using a D8 Advance X-ray diffractometer [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. The instrument was operated at 200 mA and 40 kV. The diffraction angle (2θ) was scanned from 3\u0026deg; to 40\u0026deg; with a step size of 0.04\u0026deg; and a dwell time of 0.6 seconds per step. The relative crystallinity, defined as the ratio of the crystalline peak area to the total diffraction area, was calculated using MDI Jade 6.5 software. Each sample was measured in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e\n\u003ch3\u003eFourier transform infrared spectroscopy\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe ordered structure at the starch surface was determined using a Fourier transform infrared spectrometer (FTIR, Vertex 70, Bruker, Germany) according to the protocol described [\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. The starch sample was evenly distributed on the metal mold surface of an attenuated total reflectance accessory and then exposed to an infrared beam in the range of 4000 to 400 cm⁻\u0026sup1;. Spectra were collected with 32 scans at a resolution of 4 cm⁻\u0026sup1;. The spectral data within the 1200\u0026ndash;800 cm⁻\u0026sup1; range were analyzed using OMNIC 8.2 software. Each sample was measured in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eStarch pasting properties\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe pasting properties of starch were determined using a Rapid Visco Analyzer (RVA4500, Perten, Australia). Precisely 1.96 g of starch was weighed and mixed with 26.04 g of ultrapure water to prepare a 28.00 g starch slurry at 7% (w/w). The testing profile was as follows: the sample and corresponding distilled water were first held at 50\u0026deg;C for 1 min, then heated from 50 to 95\u0026deg;C for 3.7 min, maintained at 95\u0026deg;C for 2.5 min, cooled to 50\u0026deg;C for 3.5 min, and finally held at 50\u0026deg;C for 2 min [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. Data was analyzed using the Thermal Cycle for Windows software. Each sample was measured in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eStarch thermal properties\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThermodynamic properties of starch were determined using a differential scanning calorimeter (DSC Model 200 F3 Maia, NETZSCH, Germany). Starch was mixed with distilled water at a 1:2 (g:g) ratio, sealed in an aluminum pan, and kept overnight. Using an empty sealed aluminum crucible as a reference, the temperature was increased from 20 to 100\u0026deg;C at a rate of 10\u0026deg;C min⁻\u0026sup1;. The thermodynamic parameters were recorded and calculated [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e]. After storing the samples at 4\u0026deg;C for 7 days, the retrogradation value and related parameters were measured under identical conditions. Each sample was analyzed in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eStarch iodine binding capacity, blue value, and maximum absorption wavelength\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eApproximately 80 mg of starch was weighed and added to 10 mL of a 50 mmol/L phosphate buffer (pH 7.0). The mixture was placed in a boiling water bath, shaken for 60 min, and then cooled to room temperature. 0.05 mL aliquot of the resulting solution was mixed with 4.85 mL of deionized water and 0.1 mL of iodine reagent. After rapid shaking, the mixture was allowed to stand for 15 min. Absorbance was measured using a UV-Vis spectrophotometer over a wavelength range of 500\u0026minus;700 nm. The iodine binding capacity was expressed as the ratio of absorbance at 635 nm to that at 520 nm (A635/A520) [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e]. Each sample was analyzed in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eStarch light transmittance\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe light transmittance of the starch paste were determined with slight modifications[\u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e]. starch suspension (1%, w/v) was heated in a boiling water bath for 15 min with continuous stirring. After cooling at 25\u0026deg;C for 1 h, the suspension was stored at 4\u0026deg;C for 72 hours. The light transmittance was measured at 620 nm using an ultraviolet-visible spectrophotometer (UV-1700, Shimadzu, Japan) at 0, 24, 48, and 72 h, with distilled water serving as the blank. Each sample was measured in triplicate.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eStarch solubility and swelling degree\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe solubility and swelling degree were determined with slight modifications to the described method [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e]. Precisely 160 mg starch sample (m0) was weighed into a pre-weighed 10 mL centrifuge tube (m1). Then, 8 mL of ultrapure water was added. The mixture was subjected to oscillatory heating in a water bath at 50, 60, 70, 80, and 90\u0026deg;C for 30 min, respectively. After cooling at room temperature, the sample was centrifuged at 3000\u0026times;g for 10 min, and the precipitate weight (m2) was recorded. The supernatant was transferred to another pre-weighed 10 mL centrifuge tube (m3) and evaporated to dryness. The total mass (m4) was then measured. Each sample was analyzed in triplicate. The calculation formulas are as follows:\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"No\" id=\"Taba\" border=\"1\"\u003e \u003ccolgroup cols=\"2\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSolubility (%) = (m4 / m0) \u0026times; 100%\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(1)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSwelling degree (g/g) = (m2 / m0) \u0026times; 100%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e(2)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eMean values were calculated using Excel 2021. Analysis of variance and correlation tests between datasets were performed using the Statistical Package for the Social Sciences software (version 22.0). Differences between groups were determined for significance using the Least Significant Difference method. Figures were generated using Origin 2021. Correlation analyses were performed using the R software environment.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003eSoil basic fertility\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe basic fertility of the 0\u0026minus;20 cm soil layer at in different sites are presented in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. The soil pH of HS and BD sites was 7.43, indicating a slightly alkaline condition. The pH values at TS, QHD, and CD sites ranged from 7.04 to 7.25. In terms of soil organic fertilizer, the organic matter content at the QHD site was significantly higher than at other sites, reaching 19.27 mg/kg. In contrast, the TS site had the lowest soil organic matter content of 10.09 mg/kg. The available soil nitrogen content followed the order: QHD\u0026thinsp;\u0026gt;\u0026thinsp;HS\u0026thinsp;\u0026gt;\u0026thinsp;BD\u0026thinsp;\u0026gt;\u0026thinsp;CD\u0026thinsp;\u0026gt;\u0026thinsp;TS. The QHD site exhibited a soil-available nitrogen content of 84.23 mg/kg, whereas the TS site had 54.83 mg/kg, indicating a significantly weaker nitrogen supply capacity than in other regions. Analysis of soil available phosphorus and potassium contents across different sites revealed that the HS site had the highest levels of both nutrients, with identical values of 604.67 mg/kg. Furthermore, BD, TS, QHD, and CD sites exhibited a consistent trend in soil-available phosphorus and potassium. Specifically, the BD site recorded 101.00 mg/kg; TS and QHD sites measured 137.00 mg/kg and 133.00 mg/kg, respectively, the CD site had an available nutrient level identical to that of the QHD site. In summary, the sites exhibited significant variation in soil fertility. The QHD site exhibited sufficient soil organic matter and alkali-hydrolyzable nitrogen, the HS site demonstrated significantly higher available nutrient level. In contrast, the overall soil fertility at the TS site was relatively weak. This regional heterogeneity in soil fertility provide a valuable reference for regional cultivation nutrient management of waxy maize.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eClimatic ecological factors\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe climatic ecological factors for the period from May to August across different sites are presented in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. The highest temperatures at the HS site from May to August were significantly higher than those at other sites, reaching 40.33\u0026deg;C in June. In contrast, the CD site exhibited significantly low temperatures, with the lowest in May at 0.54\u0026deg;C and in August at 2.77\u0026deg;C, its overall temperature was lower than that of the other sites. The average temperature from May to August followed the trend of HS\u0026thinsp;\u0026gt;\u0026thinsp;TS\u0026thinsp;\u0026gt;\u0026thinsp;BD\u0026thinsp;\u0026gt;\u0026thinsp;QHD\u0026thinsp;\u0026gt;\u0026thinsp;CD. At the HS site, the monthly average temperature remained above 22\u0026deg;C, whereas at the CD site, the average temperature for July and August remained below 21\u0026deg;C. Analysis of the effective accumulated temperature indicated that the total effective accumulated temperature from May to August at the HS site reached 2117.28\u0026deg;C, the highest among all sites, with monthly accumulations exceeding 500\u0026deg;C (reaching 615.02\u0026deg;C in July). However, the total effective accumulated temperature at the CD site was 925.07\u0026deg;C, which was less than half that at the HS site. The effective accumulated temperature in May was approximately 66.35\u0026deg;C, the thermal conditions at this site were significantly weaker than those in other regions. Analysis of precipitation across the sites exhibited an uneven spatial distribution. TS and QHD sites experienced concentrated rainfall in August (252.96 and 250.05 mm, respectively). At the BD site, precipitation in July and August accounted for 75% of the total. The highest rainfall values recorded at HS and CD sites were 126.74 and 248.85 mm, respectively, in July. In summary, meteorological conditions varied significantly among the five sites. The HS site had sufficient heat; however, relatively low rainfall. In contrast, the CD site was characterized by insufficient heat and a more uniform precipitation distribution. TS and QHD sites exhibited concentrated rainfall in the later growth period. This regional heterogeneity in ecological factors provides fundamental data to support research on the regional adaptability of waxy maize.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe grain nutritional quality of waxy maize in different ecological regions\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSugar\u003c/p\u003e \u003cp\u003e(mg/g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTotal starch\u003c/p\u003e \u003cp\u003e(mg/g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eAmylopectin\u003c/p\u003e \u003cp\u003e(mg/g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eAmylose\u003c/p\u003e \u003cp\u003e(mg/g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAmylopectin proportion(%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e79.58b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e429.49c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e428.60b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.89c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e99.79a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e87.14a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e456.57a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e455.65b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.92a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e99.80a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e84.21ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e458.10a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e457.27b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.83a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e99.82a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eQHD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e82.01ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e442.81b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e441.41a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1.41b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e99.68a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e73.31c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e455.32a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e454.37b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.96a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e99.79a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003eData are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec20\" class=\"Section2\"\u003e \u003ch2\u003eGrain texture properties\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eAcross the different sites, significant differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) in the texture characteristics of waxy maize kernels were observed (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Grain hardness was highest at the TS site, followed by the BD site, and lowest at the HS site. Grain cohesiveness was highest at the HS site, while BD and TS sites exhibited lower values. The grain elasticity exhibited no significant difference (P\u0026thinsp;\u0026gt;\u0026thinsp;0.05) across sites, suggesting that the experimental location did not affect it. Grain adhesiveness was significantly higher (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) at the HS site than at BD and TS sites, with no significant difference between BD and TS sites. Grain chewiness was highest at the QHD site and lowest at the HS site; the remaining sites exhibited a medium level with no significant differences among them.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe volume distribution and average granule size of waxy maize starch in different ecological regions\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eD(4,3)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eD(3,2)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003eVolume distribution (\u0026micro;m)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;5\u0026micro;m\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5\u0026ndash;15\u0026micro;m\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026gt;\u0026thinsp;15\u0026micro;m\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e12.79e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e8.33e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e9.39a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e46.48a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e44.14e\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.45d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.37c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.00c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e34.69c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e57.31d\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e22.49a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11.08a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e6.62e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e26.06e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e67.32a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eQHD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e15.00b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e9.27d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e8.57b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e33.75b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e57.69c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.81c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e10.00b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e7.06d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e32.05d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e60.88b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003eData are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec21\" class=\"Section2\"\u003e \u003ch2\u003eGrain nutritional quality\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe grain nutritional quality of waxy maize exhibited significant differences across the sites (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The soluble sugar content in the grains was significantly higher at the BD site compared to HS and CD sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). TS and QHD sites exhibited intermediate values, indicating no significant difference from the BD site, while the CD site recorded the lowest sugar content. Regarding total starch content, the TS site had the highest level, the BD and CD sites did not differ significantly from TS, Thses three sites had significantly higher total starch content than QHD and HS sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), with the HS site exhibiting the lowest value. For amylose content, the QHD site had the highest level, which was significantly greater than that at all other sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The HS site exhibited the lowest amylose content, and no significant differences were observed among BD, TS, and CD sites. The amylopectin content in the grains was significantly higher at the QHD site compared to the other sites (P\u0026thinsp;\u0026gt;\u0026thinsp;0.05).\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe pasting properties of waxy maize starch in different ecological regions\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePV(cP)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTV(cP)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFV(cP)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eBD(cP)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSB(cP)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003ePtemp(\u0026deg;C)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1441.3b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e664.7c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e772.0d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e760.0c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e107.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e77.8a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1859.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e871.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e941.7b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e988.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e70.7a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e68.7a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1824.7a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e906.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e976.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e918.7ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e70.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e77.5a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eQHD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1340.3b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e533.0d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e637.3e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e807.3bc\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e104.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e76.7a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1788.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e795.7b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e876.7c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e992.3a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e81.0a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e74.0a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003eData are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec22\" class=\"Section2\"\u003e \u003ch2\u003eStarch granule morphology, optical microscopy, and polarized cross structure\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe scanning electron microscope (SEM) plays a crucial role in characterizing the structure of starch granules. SEM images of starch from different sites revealed potential variations in the size and shape of starch granules across treatments. Starch granule size significantly influences the quality characteristics of isolated starch. The starch granules from the HS site were relatively small and uniformly distributed, with relatively smooth surfaces (Figs.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e-A-B). In contrast, the starch granules from the TS site were significantly larger than those from other sites, with nearly spherical shapes and plump surfaces. This observation aligns with the results described in Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e, which indicates that the TS site has the largest volume average particle size. Optical microscopy (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e-C) indicated that the waxy corn starch granules from the HS site were well dispersed with minimal aggregation. Conversely, the starch granules from the TS site exhibited a higher degree of aggregation, attributed to stronger surface forces associated with larger particle sizes. In polarized-light micrographs (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e-D), waxy corn starch from all sites exhibited the typical Maltese cross extinction pattern, indicating the presence of crystalline starch. The extinction cross of starch from the TS site exhibited greater clarity, consistent with its highest crystallinity characteristic as presented in Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. In contrast, the extinction cross of starch from the QHD site appeared relatively indistinct, consistent with its moderate starch crystallinity.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe thermal properties of waxy maize starch in different ecological regions\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"6\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTo(℃)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eTp(℃)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eTc(℃)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eΔH(J/g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eR(%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e70.52b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e78.53ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e84.67a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.03d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e32.22b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e70.75b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e79.33ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e85.17a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e6.31b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e24.94c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e81.30a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e83.20a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e88.76a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e7.22a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e30.21b\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eQHD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e70.69b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e73.63b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e76.62b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.62c\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e38.79a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e65.32b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e77.56ab\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e79.84b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.06d\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e22.29c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"6\"\u003eData are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec23\" class=\"Section3\"\u003e \u003ch2\u003eStarch size distribution\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThere were significant differences in the average particle size and volume distribution of waxy corn starch among different sites (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Regarding the average starch granule size, both the volume-average particle size D (4,3) and the surface-area mean diameter D (3,2) at the TS site were significantly higher than those at other sites (\u0026lt;\u0026thinsp;15 \u0026micro;m) at 44.14%. In contrast, the TS site exhibited the opposite trend: the volume proportion of large starch particles (\u0026gt;\u0026thinsp;15 \u0026micro;m) reached 67.32%, significantly higher than at other sites, while the proportion of medium starch particles (5\u0026ndash;15 \u0026micro;m) was only 26.06%. These results indicate that large particles dominated the waxy maize starch granules at the TS site, whereas those at the HS experimental site were primarily small particles. The starch particle size volume distribution curve (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e-A) indicates that the peak of the starch size distribution at the TS site shifted towards the large particle size range. In contrast, the distribution peak at the HS experimental site was concentrated in the small-particle size range. This observation aligns with the results in Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e, where the volume-average particle size was largest at the TS experimental site and smallest at the HS site, providing a visual demonstration of the differences in particle size among the sites.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec24\" class=\"Section2\"\u003e \u003ch2\u003eStarch relative crystallinity and short-range ordered structure\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe crystallinity of starch is associated with the ordered structure of amylopectin molecules within the granules. XRD patterns (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e-B) revealed that starch from all sites exhibited typical type A crystalline characteristics, with diffraction peaks at 15\u0026deg;, 17\u0026deg;, 18\u0026deg;, and 23\u0026deg;. Differences in starch relative crystallinity were observed among the sites. The TS site exhibited significantly higher relative crystallinity than the other sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), the HS site had the lowest value. The remaining sites exhibited no significant differences.In the FTIR full spectra (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e-C), the absorption peak trends in regions, including 3400 cm⁻\u0026sup1; (hydroxyl stretching vibration), 2930 cm⁻\u0026sup1; (C-H stretching vibration), and 1640 cm⁻\u0026sup1; (water molecule bending vibration), were consistent across all sites. However, differences in IR ratios were observed within the characteristic peak region (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e-D). The 1045/1022 ratio at the HS site was significantly lower than that at the other sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), the 1022/995 ratio exhibited no significant differences among the sites.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec25\" class=\"Section3\"\u003e \u003ch2\u003eStarch pasting properties\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe peak viscosity (PV) of starch from the BD and TS sites was significantly higher than that from HS and QHD sites presented in Table\u0026nbsp;\u003cspan refid=\"Tab6\" class=\"InternalRef\"\u003e6\u003c/span\u003e. The trend of trough viscosity (TV) aligned with that of final viscosity. The QHD site exhibited the lowest TV. The breakdown value reflects the thermal stability of starch. The breakdown values for BD and CD sites were significantly higher than those for the HS site (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Furthermore, no significant differences were observed in setback value and starch pasting temperature among the sites.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab6\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 6\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eThe iodine binding capacity, blue value and maximum absorption wavelength of waxy maize starch in different ecological regions.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSite\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eIodine binding capacity\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eBlue value\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eMaximum absorption wavelength(nm)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.031b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.887b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e547.00ab\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.072b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.727a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e553.75a\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eTS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.072b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.898a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e542.25bc\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eQHD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.159a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.897a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e537.00c\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.079b\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2.359a\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e540.25bc\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"4\"\u003eData are means of three replications. Means with no letter in common indicate significant differences between regulator by least significant difference test (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec26\" class=\"Section3\"\u003e \u003ch2\u003eStarch thermal properties\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe QHD site exhibited a smaller area under the starch heat flow peak(Table\u0026nbsp;7), corresponding to its lower pasting enthalpy change. This visually demonstrates the differential regulation of starch thermal pasting properties across the sites. Specifically, the ΔHgel was highest at the TS site, significantly exceeding that at HS, QHD, and CD sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). HS and CD sites recorded the lowest ΔHgel values. The retrogradation value was significantly higher at the QHD site than at the other sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05), whereas BD and CD sites exhibited lower values. This suggests that the starch at the QHD site is more susceptible to pasting.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec27\" class=\"Section3\"\u003e \u003ch2\u003eStarch iodine binding capacity, blue balue, and maximum absorption wavelength\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe starch from the QHD site exhibited significantly higher (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) iodine binding capacity than that from the other sites (Table\u0026nbsp;8). HS, BD, TS, and CD sites indicated no significant differences. This indicates that the starch from QHD possesses a stronger ability to bind iodine, which can be associated with its amylose content or molecular chain structure. This finding aligns with the result presented in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e, which indicates that the QHD site had the highest amylose content. The blue value of starch from the HS site was significantly lower than that from the other sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). BD and CD site both exhibited relatively high blue values with no significant difference between them. The blue value further reflects differences in the color development intensity of the starch-iodine complex, and its variation trend is consistent with that of the iodine binding capacity. The BD site exhibited the highest maximum absorption wavelength for starch, while the QHD site demonstrated the lowest, with the QHD site being significantly lower than the BD site (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Additionally, no significant differences were observed among the remaining sites.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec28\" class=\"Section2\"\u003e \u003ch2\u003eStarch light transmittance\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe starch light transmittance at each site decreased with prolonged storage time at 4\u0026deg;C (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e-A). The transmittance at the TS site was significantly higher than that at the other sites at all time sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). At the TS site, the starch light transmittance reached 80% at 0 and 24 h; however, it was reduced below 80% at 48 and 72 h. At the other sites, the initial transmittance values at various time sites ranged from 60% to 75%. At 0 h, the starch light transmittance at the TS site was significantly greater than that at the other sites, while no significant differences were observed among the other sites. At 72 h, the TS site remained significantly higher than HS and QHD sites; however, it was not significantly different from BD and QHD sites. The TS site exhibited the largest decline in starch light transmittance, suggesting that its starch paste had better initial transparency but weaker long-term stability.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec29\" class=\"Section2\"\u003e \u003ch2\u003eStarch solubility and swelling power\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe starch solubility at all sites increased with increasing temperature (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e-B). Solubility and swelling power exhibited a significant correlation with temperature, a phenomenon attributed to the increased mobility of starch and water molecules, with higher temperatures promoting the diffusion of amylose and amylopectin. When the temperature reached 90\u0026deg;C, the starch swelling power at the TS site was significantly higher than that at the other sites (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). This finding is associated with the starch structural characteristics of the TS site (large particle size and high crystallinity) as presented in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. On the one hand, the larger particle size provides more physical space for the starch, resulting in a greater expansion volume after water absorption. On the other hand, the hydrogen bond network in the crystalline region of starch with high crystallinity undergoes extensive dissociation at high temperatures. The strong water absorption driving force during this dissociation process allows it to bind more water. In contrast, the starches from other sites (HS and BD) have smaller mean particle sizes and lower relative crystallinity, lacking sufficient swelling space and exhibiting a weaker water-absorption driving force. Consequently, their starch swelling power was significantly lower than that of the TS site. The solubility of starch at each site increased with increasing temperature (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e-C). Within the 50\u0026ndash;90\u0026deg;C temperature range, starch solubility at all sites increased progressively with increasing temperature. A greater temperature rise corresponded to a more pronounced rate of increase in solubility. When the temperature reached 90\u0026deg;C, starch solubility at each site peaked; however, the differences were not statistically significant.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eCorrelation analysis between grain texture and starch physicochemical properties\u003c/h3\u003e\n\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003ePearson correlation coefficients and mantel test results for the relationships between grain texture and starch physicochemical properties are presented in Fig.\u0026nbsp;4-A. Starch granule size positively regulates grain texture. Stickiness and chewiness exhibited strong positive correlations with small starch granules, indicating that the tested parameters have a relatively lesser influence on cohesiveness. The associations between indicators, including Tp and Tc, and all texture characteristics were not statistically significantly, indicating that these parameters were not key factors in regulating grain texture in this study. In summary, starch granule size is a core indicator regulating grain stickiness, chewiness, and hardness, R% primarily influences grain elasticity. These results provide a quantitative basis for targeted improvement in grain texture quality by optimizing starch physicochemical properties.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003cdiv id=\"Sec31\" class=\"Section2\"\u003e \u003ch2\u003ePrincipal component analysis of soil conditions, meteorological factors, and grain quality\u003c/h2\u003e \u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe results of principal component analysis (Fig.\u0026nbsp;4-B) indicated that the first two principal components (PC) cumulatively explained 58.8% of the total variation in grain quality-associated indicators of waxy maize across different ecological regions in northern China. PC1, accounting for 34.9% of the variance, represented the dimension of structural strength and thermal stability, its positive direction was significantly associated with variables, including the proportion of large-sized starch (\u0026gt;\u0026thinsp;15 \u0026micro;m), relative crystallinity of starch, peak gelatinization viscosity, grain hardness, and soil available nitrogen content. Conversely, the negative direction was associated with variables including the proportions of small sized starch (\u0026lt;\u0026thinsp;5 \u0026micro;m and 5\u0026ndash;15 \u0026micro;m), adhesiveness, chewiness, rainfall, and retrogradation value. PC2, explaining 23.9% of the variance, corresponded to the dimension of starch composition and pasting properties, its positive direction was associated with high amylose content, a strong setback tendency, and a relatively high pasting temperature.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThis study indicates that differences in grain texture of waxy maize across various ecological regions are primarily due to the regulatory effects of environmental factors on starch granule morphology and crystal structure. Specifically, the TS site exhibited the highest kernel hardness, whereas the HS site demonstrated the lowest hardness and relatively high adhesiveness. This phenomenon is closely associated with the starch granule size distribution, the TS site was dominated by large-granule starch (\u0026gt;\u0026thinsp;15 \u0026micro;m), while the HS site was rich in small-granule starch (\u0026lt;\u0026thinsp;5 \u0026micro;m). Multivariate correlation analysis (Fig.\u0026nbsp;6) further revealed that the proportion of small granule starch (\u0026lt;\u0026thinsp;5 \u0026micro;m) exhibited a significant positive correlation (P\u0026thinsp;\u0026le;\u0026thinsp;0.01) with both adhesiveness and chewiness of the kernels. The underlying mechanism is associated with the greater specific surface area of small granule starch, which promotes the interactions between granules and molecules, thereby contributing to a more pronounced sticky taste during mastication. This observation aligns with the conclusion that increasing the proportion of small granule starch improves the viscoelasticity of food products [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. Conversely, large granule starch can provide a more supportive physical structure [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. The TS site not only exhibited a larger starch granule size but also the highest relative crystallinity. The synergistic effect of these two factors formed a dense crystalline network, significantly increasing the mechanical hardness of the kernels. This finding is consistent with the established theory that both starch crystallinity and granule size collectively determine hardness [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]. However, the HS site was characterized by a predominance of small granule starch and low crystallinity, resulting in loose granule binding and, consequently, reduced hardness [\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe crystalline structure of starch is a critical factor that interacts with environmental factors to determine its final functional properties. XRD and FTIR analyses consistently indicate that starch from the TS site exhibits the highest crystallinity, while that from the HS site demonstrates the lowest. This variation correlates positively with particle size, as larger granules provide more space for the orderly arrangement of starch molecular chains, which facilitates the formation and maintenance of well-defined crystalline regions [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. In contrast, the crystalline structure of small granule starch tends to be looser and less complete [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]. The lower 1045/1022 cm⁻\u0026sup1; FTIR peak intensity ratio observed for the HS site further confirms its insufficient crystalline order. These differences in crystallinity directly determine the thermodynamic behavior of the starch. ΔH represents the energy required to disrupt the double helix structure within the starch crystalline region [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. The high crystallinity of starch from the TS site suggests a denser, more stable internal hydrogen-bond network, thereby requiring greater energy input for gelatinization, which aligns with findings reported in rice studies [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]. The higher PV values observed at TS and BD sites are due to the starches at these sites, which possess both large granules and high crystallinity. Large granule starch exhibits a stronger water absorption and swelling potential during the initial stage of gelatinization, while a highly crystalline structure offers greater resistance to heat shear. The synergistic effect of these two factors [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e] allows starch to accumulate and reach a higher viscosity peak during heating [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eEnvironmental conditions modify the chemical composition and molecular structure of starch by regulating the starch biosynthesis pathway, thereby affecting its functional properties. In this study, the TS site exhibited the highest total starch content, likely because its favorable environment effectively promoted starch synthase activity and substance accumulation [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e]. Conversely, waxy maize at the HS site exhibited the lowest total starch content, which can result from suppression of starch synthesis gene expression under environmental stress. Notably, the total starch content of waxy maize at the QHD site was not the highest; however, its amylose content was significantly higher than at other sites, resulting in a substantial increase in setback viscosity. This is attributed to the greater tendency of amylose molecules to rearrange and form a hard gel during the cooling process [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. Furthermore, the strongest iodine-binding capacity of waxy maize starch from the QHD site directly confirmed its higher amylose content [\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]. The lower starch blue value at the HS site reflected its lower amylose content. Differences in the λmax of starch across the sites suggested that environmental conditions could influence fine molecular structures, including the amylopectin branch-chain length. The starch granule and crystal structure indirectly determine its performance after gelatinization. The highest initial transmittance of waxy maize starch from the TS site could be attributed to a more uniform gel network formed after gelatinization of its large, highly crystalline granules. Waxy maize at the TS site exhibited the highest starch swelling power at 90\u0026deg;C because the high temperature sufficiently disrupted its dense crystalline structure, resulting in significant dissociation of hydrogen bonds, while the large granules provided additional space to bind more water [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e]. Furthermore, the higher kernel soluble sugar content observed at the BD site can be attributed to lower efficiency in converting photosynthetic products into starch under its specific environmental conditions, including temperature and moisture, resulting in sugar accumulation. This indicates environmental plasticity in the distribution of carbon metabolic fluxes.\u003c/p\u003e \u003cp\u003eMeteorological factors exhibited significant spatial heterogeneity in shaping the quality of waxy maize kernels. In regions represented by the HS site, the grain-filling stage (May to August) was characterized by abundant heat resources, with an effective accumulated temperature of 2117.28\u0026deg;C; however, relatively low precipitation of 333.90 mm. The favorable thermal conditions likely accelerated starch biosynthesis, promoted the formation of small starch granules, and optimized grain viscoelasticity. In contrast, the CD site experienced a lower effective accumulated temperature of 925.07\u0026deg;C during grain filling, with evenly distributed rainfall. However, the low temperatures likely inhibited starch synthase activity, resulting in insufficient starch granule development and, consequently, reduced grain adhesiveness and chewiness. These findings align with existing research, indicating that high-temperature treatment induces irregular surface morphology and increases the average starch granule size of waxy maize starch, while reducing its gelatinization viscosity and enthalpy value [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. Conversely, moderate low temperature decreases the average starch granule size [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e], increases the proportion of short chains in amylopectin, and reduces relative crystallinity, ultimately impacting grain stickiness. Similar studies in rice have reported that sustained high temperatures modify starch component accumulation, particle morphology, and crystal structure, negatively impacting edible quality [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e]. Conversely, low-temperature stress increases amylose content and retrogradation value and modifies the fine structure of amylopectin [\u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. Furthermore, TS and QHD sites experienced heavy rainfall (\u0026gt;\u0026thinsp;250 mm) during the late grain-filling stage (August), which induced grain metabolic water stress. This disturbance could have disrupted the normal construction and stability of starch structure, thereby affecting the final texture. This observation is consistent with previous reports that waterlogging stress reduces starch content and modifies its gelatinization properties [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eSoil conditions are another key driver of regional differentiation in waxy maize quality. In this study, the soil at the QHD site was rich in organic matter and readily available nitrogen. Therefore, the grains exhibited significantly higher amylose and amylopectin contents, as well as a higher iodine-binding capacity. However, the clarity of the starch polarization cross was poorer, and gelatinization stability was weaker, sufficient nitrogen supply can promote starch synthesis, and thus affect starch properties to a certain extent [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]. The soil at the HS site had higher levels of readily available phosphorus and potassium, indicating a significant nutrient supply potential. However, the grains exhibited the lowest total starch content, relative crystallinity, and swelling power. The starch granules were predominantly small, smooth, and well dispersed. This suggests that elevated soil phosphorus and potassium levels can have inhibited the full development and orderly aggregation of starch granules [\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]. At the TS site, the soil had low organic matter content and poor nitrogen supply capacity.The starch exhibited the largest average granule size, the highest crystallinity and enthalpy value, as well as excellent thermal stability of gelatinization. Additionally, the grain hardness was relatively higher. This reveals an interesting phenomenon, low nitrogen soil conditions promote the development of waxy maize starch granules toward larger size and higher crystallinity, forming a more compact starch granule structure [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]. Soil fertility at the BD site was medium. The grains exhibited higher kernel soluble sugar content, PV, and breakdown value; however, the starch's thermal stability was only moderate. Previous studies have suggested that sandy soil has a poor water-retention capacity under drought conditions, thereby exacerbating the inhibitory effect of water stress on starch synthesis. In contrast, the stronger water-retention capacity of clay soil can buffer the adverse effects of excessive water on the stability of the starch structure under rainy conditions [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e]. This provides a theoretical basis for explaining the quality differences among different soil types within the same climatic region.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003e \u003cdiv class=\"BlockQuote\"\u003e \u003cp\u003eThe regional heterogeneity of meteorological, ecological, and soil factors constitutes the core determinant of waxy maize grain quality and starch properties. The hydrothermal combination characteristics in different ecological regions exert significantly distinct regulatory effects on starch granule development, crystalline structure, and functional properties. At the HS site in the central-southern Hebei region, abundant heat supply coupled with precipitation scarcity during the growth period shaped starch characteristics, resulting in small particle size and low crystallinity, conferring a quality advantage in higher grain adhesiveness. In BD, concentrated rainfall during the later growth stage resulted in starch granules of medium size and medium crystallinity. This resulted in higher grain hardness and soluble sugar content, albeit with weaker starch thermal stability. In the northern Hebei region, insufficient heat supply at the CD site inhibited the orderly development of starch granules, directly resulting in overall quality deterioration in waxy maize. The regulatory effects of concentrated precipitation during the later growth stage differed significantly between the two experiment sites in the Eastern Hebei region. At the TS site, a starch crystal structure characterized by large particle size and high crystallinity formed, resulting in synergistic increases in grain hardness, total starch accumulation, and starch PV. In contrast, the QHD site exhibited a higher proportion of amylose in the grains and improved iodine-binding capacity, thereby strengthening the starch retrogradation characteristics. Therefore, the waxy maize from this site exhibited unique aging resistance and gel functional properties. This study clarifies the key influencing factors and regulatory mechanisms underlying quality formation in waxy maize across different ecological regions of Hebei Province. This study provide precise theoretical support for the regional cultivation of high-quality waxy maize and the breeding of special varieties.\u003c/p\u003e \u003c/div\u003e \u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eFunding\u003c/h2\u003e \u003cp\u003eThis study was financially supported by the National Key Research and Development Program of China (Grant No.2023YFD2301505).\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003ePegntao Ji: investigation, data curation, writing\u0026mdash;original draft. Xiangling Li: investigation,methodology,writing\u0026mdash;original draft. Weixin Dong: methodology, visualization. Peijun Tao: resources, writing\u0026mdash;review and editing. Yuechen Zhang: Funding acquisition, writing\u0026mdash;review and editing. All authors have read and agreed to the published version of the manuscript.\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eData will be made available on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eLu, D. L. \u0026amp; Lu, W. P. Effects of protein removal on the physicochemical properties of waxy maize flours. \u003cem\u003eStarch-St\u0026auml;rke\u003c/em\u003e \u003cb\u003e64\u003c/b\u003e, 874\u0026ndash;881. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1002/star.201200038\u003c/span\u003e\u003cspan address=\"10.1002/star.201200038\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2012).\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYu, X. R. et al. Endosperm structure and physicochemical properties of starches from normal, waxy, and super-sweet maize. \u003cem\u003eInt. J. 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Agr\u003c/em\u003e. \u003cb\u003e15\u003c/b\u003e, 309\u0026ndash;316. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.ijbiomac.2019.02.018\u003c/span\u003e\u003cspan address=\"10.1016/j.ijbiomac.2019.02.018\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e (2016).\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Waxy maize, starch, ecological region","lastPublishedDoi":"10.21203/rs.3.rs-9183115/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-9183115/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe growth environment is a critical regulatory factor in starch macromolecule biosynthesis. Variations in these natural ecological conditions can result in differences in quality within the same variety grown in different regions. The grain texture and starch physicochemical properties were examined across five different sites in Hebei Province, including Hengshui (HS), Baoding (BD), Tangshan (TS), Qinhuangdao (QHD), and Chengde (CD). Significant divergence of grain quality existed across the five sites. The grain hardness was highest at the TS site, while the adhesiveness was optimal at the HS site. The BD site exhibited the highest soluble sugar content, and the TS site had the highest total starch content. Notably, the amylopectin content at the QHD site was significantly higher than at the other locations. Regarding starch structure, the starch granules from the TS site were the largest and demonstrated the highest relative crystallinity. In contrast, the starch granules from the HS site were smaller and more uniformly distributed. Analysis of starch functional properties revealed that the peak viscosity of starch was significantly higher at BD and TS sites. The enthalpy value and swelling power of starch were optimal at the TS site, whereas the HS site exhibited higher starch light transmittance but weaker stability. Further analysis indicated that these differences were significantly associated with the soil conditions and climatic factors at each site. Overall, this study provide a theoretical basis for the application of waxy maize starch in the food industry and for improving starch quality through breeding.\u003c/p\u003e","manuscriptTitle":"Variations in grain textural and starch physicochemical properties of waxy maize across contrasting ecological regions","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-04-06 14:45:05","doi":"10.21203/rs.3.rs-9183115/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-04-20T10:16:38+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-18T09:43:24+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-16T07:40:41+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"129918789878463472377054113533582083441","date":"2026-04-08T04:46:28+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"130725082530437873106662917494395738570","date":"2026-04-07T15:36:05+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-06T04:35:08+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-04-02T09:20:57+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"273909271716107894587421224829045604171","date":"2026-04-01T05:28:42+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"193417691695139867641244740830797148420","date":"2026-04-01T03:39:53+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-04-01T03:24:03+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2026-04-01T03:17:10+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2026-03-31T12:44:22+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2026-03-25T02:56:48+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2026-03-25T02:50:59+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"511e577e-7af6-480e-87a3-aee70ece3bb0","owner":[],"postedDate":"April 6th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[{"id":65654680,"name":"Biological sciences/Biochemistry"},{"id":65654681,"name":"Biological sciences/Plant sciences"}],"tags":[],"updatedAt":"2026-05-12T06:25:22+00:00","versionOfRecord":[],"versionCreatedAt":"2026-04-06 14:45:05","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-9183115","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-9183115","identity":"rs-9183115","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}