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PLOS One logoLink to PLOS One
. 2015 Apr 13;10(4):e0123330. doi: 10.1371/journal.pone.0123330

Effects of Delaying Transplanting on Agronomic Traits and Grain Yield of Rice under Mechanical Transplantation Pattern

Qihua Liu 1,*, Xiu Wu 1, Jiaqing Ma 1,*, Bocong Chen 1, Caiyun Xin 1
Editor: Paul C Struik2
PMCID: PMC4395310  PMID: 25875607

Abstract

A delay in the mechanical transplantation (MT) of rice seedlings frequently occurs in Huanghuai wheat-rice rotation cropping districts of China, due to the late harvest of wheat, the poor weather conditions and the insufficiency of transplanters, missing the optimum transplanting time and causing seedlings to age. To identify how delaying transplanting rice affects the agronomic characteristics including the growth duration, photosynthetic productivity and dry matter remobilization efficiency and the grain yield under mechanical transplanting pattern, an experiment with a split-plot design was conducted over two consecutive years. The main plot includes two types of cultivation: mechanical transplanting and artificial transplanting (AT). The subplot comprises four japonica rice cultivars. The results indicate that the rice jointing, booting, heading and maturity stages were postponed under MT when using AT as a control. The tiller occurrence number, dry matter weight per tiller, accumulative dry matter for the population, leaf area index, crop growth rate, photosynthetic potential, and dry matter remobilization efficiency of the leaf under MT significantly decreased compared to those under AT. In contrast, the reduction rate of the leaf area during the heading-maturity stage was markedly enhanced under MT. The numbers of effective panicles and filled grains per panicle and the grain yield significantly decreased under MT. A significant correlation was observed between the dry matter production, remobilization and distribution characteristics and the grain yield. We infer that, as with rice from old seedlings, the decrease in the tiller occurrence, the photosynthetic productivity and the assimilate remobilization efficiency may be important agronomic traits that are responsible for the reduced grain yield under MT.

Introduction

In China, rice is a staple food, feeding more than 65% of the population. The Huanghuai district is one of the most important rice-production districts contributing to the high grain yield in China. In this region, the rice-wheat rotation is a dominant cropping system with japonica rice as rotation crop after the harvest of wheat, producing approximately 7500 kg of rice grain yield per hectare on average [1]. The conventional cultivation pattern of rice in this region is artificial transplanting, i.e., transplanting rice seedlings into the paddy field by hand, which is very laborious. This cultivation pattern is not only cumbersome but also labor-intensive for farmers, involving working and moving in a stooping posture in a paddy field [2]. In recent years, rural labor has become increasingly insufficient and expensive, mainly due to the migration of labor from the rural areas to the cities. Rural employment decreased from 355 million in 1995 to 314 million in 2007 in China, but the labor ratio of off-farm employment to total employment increased from 33% in 1986 to 63% in 2008 in the Shandong province of China [3,4], triggering enhanced labor costs in agricultural production [5]. The same scenario has also appeared in other developing countries, such as India, Peru, and Bangladesh [6,7,8]. Now, the artificial transplanting mode that cannot meet the requirement of rice production has been gradually replaced due to the serious deficiency in labor employment. Therefore, there is an urgent need to adopt a simple and convenient planting pattern as a substitute to resolve the problem that resulted from labor scarcity and expensive production costs.

The rice mechanical transplanting pattern, which was initially developed in Japan, has been a feasible cultivation pattern and is widely applied in Japan, Korea and other countries due to its ease of cultivation and considerable economic benefits brought by labor cost savings [9,10,11,12]. In China, rice mechanical transplanting technology has been explored since the 1950s. However, until now, the technology has not been widely applied in rice production due to relatively stagnated research based on rice production districts belonging to different cropping systems. In the rice-wheat rotation cropping system, one prominent issue preventing the application of mechanical transplantation technology is the paradox between the optimum rice seedling age at transplanting as required by the transplanting machine and the prolonged seedling age as triggered by some restricting factors [13,14,15]. The optimum rice seedling age, being conducive to the occurrence of tiller, is capable of laying a solid foundation to obtain a high grain yield. In general, young seedlings from 15–20 days of age, being characterized by suitable plant height ranging from 12–17 cm, and stubborn individual plants with enough nutrient supply from the seedling-nursery tray are suitable for mechanical transplanting [16]. These plants take less time to resume normal growth after mechanical harm to roots from being transplanted.

In the Huanghuai district of China, transplanting rice seedlings by machine is frequently delayed, even causing the seedling age to be greater than 40 days, due to the late harvest of wheat, the appearance of unfavorable weather and the limited number of transplanting machines. If sowing is delayed, some varieties with long-growth duration fail to mature normally, thereby inducing the frequent occurrence of the postponement of transplantation. Usually, rice seeds were sown in hard trays at a high density for MT. Because of intensive nutrition competition among individuals, rice seedlings, which grow for a longer duration in hard trays and are not transplanted by machine in time, could receive detrimental influences [13]. These seedlings are customarily called as “old seedlings”. Until now, the performances of agronomy and yield traits in rice from old seedlings under MT remain unclear. Therefore, there is a need to conduct related research with rice plants that are transplanted by hand as controls.

The objective of the study was to test the effects of delaying transplanting on agronomic traits that are related to the grain yield and the yield component of rice under MT. These results will provide useful references for the cultivation of rice via mechanical transplantation in rice-growing districts.

Materials and Methods

Ethics statement

We obtained the relevant permission from the corresponding institute (Shandong Rice Research Institute) for planting our materials in the field. This work was unrelated to ethics issues, and no specific permissions were required for the described field study (no specific permissions were required for these locations/activities). We confirm that the field study did not involve endangered or protected species.

Experimental site and cultivar description

The field experiments were conducted at Yutai (35°00’ N, 116°39’ E) in Shandong province, China, from May to October during 2012 and 2013. The site is located in the Huanghuai district of China, with a cultivation pattern belonging to the typical rice-wheat cropping system. The data of daily average air temperature and rainfall during the rice growth season are shown in Fig 1. In 2012, the means of the daily average air temperature and precipitation were 23.94 °C and 29.09 mm, respectively. In 2013, these values were 24.87 °C and 39.83 mm, respectively. The average temperature in 2013 was approximately 1 °C higher than that in 2012. The total precipitation during the rice growth season in 2012 was 4741 mm, being 1512 mm less than that in 2013. The soil type was clay loam, and the basic fertilities were depicted as follows: organic matter 34.00 g kg-1, total nitrogen 1.94 g kg-1, available N 74.73 mg kg-1, available P 43.98 mg kg-1 and available K 199.50 mg kg-1. Four conventional japonica rice cultivars that are widely cultivated in local production, Shengdao2572, Daliang202, Huaidao11 and Jindao263, were used in the experiment.

Fig 1. Mean daily air temperature and precipitation during rice growth in 2012 and 2013.

Fig 1

Experimental design and cultivation management

The experiment was a split-plot design with three replications. The transplanting patterns, artificial transplantation (AT) and mechanical transplantation (MT), were assigned as main plots, with four cultivars being randomly allotted to sub-plots. To maintain independent water and fertilizer managements for the two transplanting patterns, the main plots were separated by a 50-cm-wide ridge with plastic film inserted into the soil at a depth of 30 cm.

According to the local rice production, rice seeds were sown on 15 May and 18 May for the artificial and mechanical transplanting patterns, respectively. The ratio of rice seedling bed area to paddy field area was 1:20 under AT while it was 1:80 under MT. The rice seedling nursery bed dimension for each cultivar was 4.5 m2 under AT and 1.125 m2 under MT. The sowing density was 74.07 g/m2 under AT and 411.52 g/m2 under MT. The rice seeds were sown on nursery bed for seedling nursery under AT. Urea was broadcast at 7.5 g/m2 and 14.9 g/m2 when the leaf age reached 2.1 and 4. The soil of nursery bed maintained moisture before the 3-leaf stage and a shallow-water was kept after the 3-leaf stage. The nursery measurements of the rice seedlings for MT were established according to the method of Yu [16] with melioration. Briefly, emerged seeds (150 g) were sown on each plastic tray (the cavity size was 58 cm long by 28 cm wide by 2.8 cm high) containing 4 kg of soil. All of the plastic trays were placed on the nursery bed when the height of the seedlings reached approximately 1.5 cm. Before the leaf age reached 1.1, the seedlings were irrigated and then drained when the water depth was as high as the nursery bed. Subsequently, intermittent irrigation (drying-wetting alternation) was performed, i.e., the seedlings were irrigated until the surface soil in each plastic tray became dry. Urea was applied at 7.5 g/m2 when the seedling leaf age was 1.1. The rice seedlings were uniformly transplanted to the paddy field on 28 June for both of the transplanting patterns across the two study years.

When transplanted, the rice seedlings for the mechanical transplantation pattern were 42 days old with an average leaf age of approximately 4.5 and a plant height within 19 cm. A transplanter (KUBATO, SPU-60) that was made in Japan was used to complete the mechanical transplantation. The distances between the rows and hills were 30 cm and 12 cm, respectively, with 3–4 and 4–5 seedlings per hill for the artificial and mechanical transplantation patterns. The size of each sub-plot was 30 m2. Fertilizer and other agronomy managements were in accordance with local practices. The total N application amount was 270 kg ha-1 with an N, P and K application rate of 2:1:1.

Sampling and determination

After the rice seedlings were transplanted to the field, the numbers of tiller occurrences of 20 fixed hills from each plot were surveyed at 4-d intervals from 16 to 40 days in 2012 and at 5-d intervals from 11 to 51 days in 2013. The dates of rice jointing, booting, heading and maturity stages in every treatment were recorded, respectively. During the four growth stages that were mentioned above, the above-ground parts of the rice plants from 6 hills from each treatment were sampled and taken to the laboratory, where they were dried at 105 °C for 0.5 h and subsequently divided into two (leaf, and stem and sheath at the jointing stage) or three (leaf; stem and sheath; and panicle at the booting, heading and maturity stages) parts. Then, the separated organs were dried at 70 °C until constant weight and weighed to determine the dry matter weight.

At maturity, thirty plants of each plot were randomly selected to measure the plant height in the field, and the grain yield from each plot was obtained by reaping 3 m2 of rice plants (excluding the border plants). The rice grain yield was determined with the moisture content being adjusted to 14%. The agronomic traits of the panicle and yield components, including the panicle length; the number of the primary and secondary branches and the total and filled grains per panicle; the effective number of panicles per m2; and the 1000-grain weight, were measured by randomly selecting 30 rice plants (except for the border plants) from each plot.

Data processing

Based on the parameters that were explicated above, the dry matter remobilization amount, the remobilization efficiency and conversion rate, the reduction rate of the leaf area, the photosynthetic potential and the population growth rate were calculated according to the following equations [17,18]:

Dry matter remobilization amount (g) = Dry matter weight per tiller at heading-dry matter weight per tiller at maturity

Dry matter remobilization efficiency (%) = 100×dry matter remobilization amount/ dry matter weight per tiller at maturity

Dry matter conversion rate (%) = 100×dry matter remobilization amount/dry weight of the grains per tiller at maturity

Leaf area index (LAI) = leaf area of the land area per m2

Reduction rate of leaf area (m2 d-1) = (LAI2-LAI1) / (t2-t1)

Where LAI1 is the initial LAI, LAI2 is the LAI that was measured the second time, t1 and t2 represent the dates of the first and the second measurements, respectively, and the difference between these dates is the time interval of the two measurements.

Photosynthetic potential of population (m2 d) = 1/2(L1+L2) × (t2-t1)

Where L1 and L2 are the leaf areas that were measured at the first and second times, t1 and t2 represent the dates of the first and the second measurements, respectively, and the difference between these dates is the time interval of the two measurements.

Crop growth rate (g m-2 d-1) = (w2-w1) / (t2-t1)

Where w1 and w2 represent the dry matter weight of the population tested as measured the first and second times, respectively, and the difference between t2 and t1 is the time interval of the two measurements.

Statistical analysis

The data were analyzed by an analysis of variance (ANOVA) using SPSS 11.0 (Statistical Package of the Social Science). To evaluate the differences between the treatments, the means were separated by Least Significance Difference (LSD) at the 5% significance level. The cultivars and cultivation patterns were regarded as fixed factors. The graphics were generated using SigmaPlot 10.0.

Results

Performance of the main growth duration for rice from old seedlings under MT

The dates for the main growth stages of the rice under AT and MT are shown in Table 1. In 2012, compared to those under AT, the rice joining, booting, heading and maturity stages were delayed 9–13 d, 7–14 d, 6–11 d and 6–8 d, respectively, under MT. In 2013, these stages were postponed for 3–8 d, 4–5 d, 3–5 d and 4–5 d, respectively. The difference was more pronounced in 2012 than in 2013, with the delayed days for the former being 2–9 d more than those of the latter.

Table 1. Dates of the different growth stages of rice under AT and MT in 2012 and 2013.

Dates for different growth stages
Year Cultivar Cultivation pattern Jointing Booting Heading Maturity
2012 Shengdao2572 AT 2-Aug 12-Aug 21-Aug 6-Oct
MT 13-Aug 23-Aug 30-Aug 12-Oct
Daliang202 AT 4-Aug 16-Aug 25-Aug 13-Oct
MT 17-Aug 30-Aug 6-Sep 20-Oct
Huaidao11 AT 9-Aug 22-Aug 1-Sep 15-Oct
MT 19-Aug 31-Aug 7-Sep 23-Oct
Jindao263 AT 10-Aug 24-Aug 2-Sep 17-Oct
MT 19-Aug 1-Sep 8-Sep 24-Oct
2013 Shengdao2572 AT 30-Jul 8-Aug 17-Aug 3-Oct
MT 3-Aug 13-Aug 22-Aug 7-Oct
Daliang202 AT 4-Aug 14-Aug 22-Aug 8-Oct
MT 7-Aug 18-Aug 27-Aug 13-Oct
Huaidao11 AT 5-Aug 16-Aug 26-Aug 12-Oct
MT 13-Aug 20-Aug 29-Aug 16-Oct
Jindao263 AT 6-Aug 17-Aug 26-Aug 13-Oct
MT 10-Aug 22-Aug 31-Aug 18-Oct

Dynamics of tiller occurrence for rice from old seedlings under MT

The dynamics of tiller occurrence under AT and MT are shown in Fig 2. The tiller number for all of the examined cultivars under MT exhibited a continuous downward trend compared to that under AT in 2012 and 2013. In 2012, compared to that under AT, the tiller numbers under MT decreased by 43.22%, 27.00%, 24.06%, and 35.78%, respectively, in Shengdao2572, Daliang202, Huaidao11 and Jindao263. These values decreased by 20.05%, 39.53%, 23.07%, and 18.80%, respectively, in 2013.

Fig 2. Dynamics of the tiller occurrence under AT and MT in 2012 and 2013. * = p≤0.05 and NS = nonsignificant.

Fig 2

Above-ground dry matter accumulation traits for rice from old seedlings under MT

As shown in Table 2, the rice dry matter weights (leaf; stem and sheath; and leaf + stem + sheath) at the jointing and booting stages under MT were significantly higher than were those under AT across the two study years, with every variety exhibiting the same change trend. Compared to that under AT, the dry matter weight of the leaf; stem and sheath; and the total above-ground parts at the jointing stage under MT decreased by 31.28%, 17.59% and 25.70% in 2012 and 29.35%, 15.02% and 23.33% in 2013, respectively. Similarly, the corresponding parameters at the booting stage decreased by 24.10%, 18.07% and 21.00% in 2012 and 31.23%, 20.63% and 26.06% in 2013, respectively. In addition, the rice dry matter weights (leaf, stem and sheath, and leaf + stem + sheath + panicle) except for that of the panicle during the heading stages under MT were significantly greater than were those under AT in both years. Unlike the performance of the rice during the above three stages, the dry matter weights of the panicle and the total above-ground parts at maturity exhibited different change tendencies under MT across the two study years. There were obvious varietal differences in the dry matter weight of the leaf, stem and sheath, and panicle, as well as the total above-ground parts, at maturity under MT. In 2012, the difference in the dry matter weight of the leaf between AT and MT was insignificant except for the Shengdao2572 variety. In addition, the dry matter weight of the stem and sheath for Shengdao2572 and Daliang202 significantly decreased, while that of the other varieties did not exhibit a significant change under MT. In 2013, the dry matter weight of the leaf for Daliang202, Huaidao11 and Jindao263 significantly increased under MT, but the opposite occurred for Shengdao2572. For most of the varieties, the dry matter weight of the stem and sheath under MT was not significantly altered excluding Shengdao2572; The dry matter weights of the panicles and the total above-ground parts significantly decreased by 36.79% and 25.69% in 2012 and 29.85% and 19.22% in 2013, respectively.

Table 2. Above-ground-part dry matter weight per tiller in rice at the jointing, booting, heading and maturity stages under AT and MT in 2012 and 2013.

Dry weight at jointing (g tiller-1) Dry weight at booting (g tiller-1) Dry weight at heading (g tiller-1) Dry weight at maturity (g tiller-1)
Year Cultivar Cultivation pattern Leaf Stem and sheath Total Leaf Stem and sheath Total Leaf Stem and sheath Panicle Total Leaf Stem and sheath Panicle Total
2012 Shengdao2572 AT 0.510a 0.410a 0.920a 0.786a 0.871a 1.656a 0.778a 0.925a 0.395a 2.098a 0.436a 0.868a 1.919a 3.222a
MT 0.344b 0.344b 0.687b 0.502b 0.588b 1.091b 0.482b 0.719b 0.263b 1.464b 0.346b 0.703b 1.117b 2.165b
Daliang202 AT 0.469a 0.380a 0.849a 0.879a 0.759a 1.638a 0.886a 1.064a 0.253a 2.202a 0.525a 0.939a 2.013a 3.478a
MT 0.415b 0.282b 0.697b 0.645b 0.561b 1.206b 0.694b 0.879b 0.319a 1.893b 0.521a 0.806b 1.241b 2.568b
Huaidao11 AT 0.636a 0.427a 1.063a 0.850a 0.928a 1.778a 0.884a 1.089a 0.370a 2.343a 0.562a 0.965a 2.332a 3.859a
MT 0.454b 0.386b 0.840b 0.705b 0.864b 1.569b 0.657b 0.984b 0.359a 2.000b 0.547a 0.912a 1.575b 3.034b
Jindao263 AT 0.821a 0.466a 1.288a 0.767a 0.862a 1.629a 0.804a 1.234a 0.361a 2.399a 0.484a 1.102a 2.489a 4.076a
MT 0.461b 0.375b 0.837b 0.639b 0.789b 1.428b 0.728b 1.099b 0.356a 2.182b 0.440a 1.067a 1.600b 3.108b
2013 Shengdao2572 AT 0.603a 0.496a 1.099a 0.703a 0.681a 1.384a 0.677a 0.833a 0.366a 1.876a 0.386a 0.784a 1.685a 2.855a
MT 0.386b 0.474b 0.861b 0.426b 0.448b 0.874b 0.353b 0.506b 0.182b 1.040b 0.259b 0.495b 0.899b 1.652b
Daliang202 AT 0.575a 0.448a 1.023a 0.831a 0.666a 1.497a 0.792a 0.938a 0.293a 2.023a 0.348b 0.825a 1.799a 2.971a
MT 0.375b 0.316b 0.690b 0.545b 0.536b 1.081b 0.691b 0.873b 0.326a 1.891b 0.432a 0.793a 1.229b 2.454b
Huaidao11 AT 0.535a 0.372a 0.907a 0.772a 0.747a 1.518a 0.677a 0.985a 0.296a 1.958a 0.431b 0.879a 2.135a 3.444a
MT 0.471b 0.348b 0.819b 0.523b 0.626b 1.149b 0.613b 0.921b 0.319a 1.853b 0.532a 0.884a 1.721b 3.138b
Jindao263 AT 0.662a 0.402a 1.064a 0.867a 0.921a 1.788a 0.683a 0.950a 0.270a 1.903a 0.436b 0.885a 2.180a 3.501a
MT 0.446b 0.322b 0.768b 0.688b 0.783b 1.471b 0.616b 0.873b 0.295a 1.784b 0.593a 0.858a 1.622b 3.073b

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

For every variety, the dry matter accumulation amounts of the rice population under MT were significantly lower than were those under AT across the two study years. Compared to those under AT, the dry matter accumulation amounts of the rice population under MT significantly decreased by 29.62% and 40.27% from the jointing to the booting stage, 50.69% and 35.42% from the booting to the heading stage, and 47.59% and 37.11% from the heading to the maturity stage, respectively, in 2012 and 2013 (Table 3).

Table 3. Dry matter accumulative amount of rice population based on three growth stages (from jointing to booting, booting to heading and heading to maturity) under AT and MT in 2012 and 2013.

From jointing to booting stage From booting to heading stage From heading to maturity
Year Cultivar Cultivation pattern Dry matter accumulation (t hm-2) Dry matter accumulation (t hm-2) Dry matter accumulation (t hm-2)
2012 Shengdao2572 AT 1.807a 2.426a 4.202a
MT 0.971b 0.745b 2.108b
Daliang202 AT 1.276a 2.842a 4.334a
MT 0.674b 1.651b 2.834b
Huaidao11 AT 3.449a 3.537a 5.237a
MT 2.984b 1.213b 2.672b
Jindao263 AT 3.524a 4.317a 6.425a
MT 2.448b 2.861b 2.972b
2013 Shengdao2572 AT 2.292a 1.507a 3.742a
MT 0.614b 0.299b 2.079b
Daliang202 AT 2.075a 2.596a 3.956a
MT 1.141b 2.108b 2.667b
Huaidao11 AT 2.448a 2.666a 4.130a
MT 1.475b 2.068b 2.620b
Jindao263 AT 5.054a 1.051a 3.654a
MT 3.860b 0.575b 2.371b

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

Above-ground dry matter remobilization and distribution characteristics for rice from old seedlings under MT

Compared to those under AT, the leaf dry matter remobilization amount, efficiency and contribution to the grain yield under MT significantly decreased by 55.66%, 38.60% and 29.46% in 2012 and 53.63%, 44.23% and 31.43% in 2013, respectively. The dry matter remobilization amount and the efficiency of the stem and sheath under MT markedly decreased by 55.85% and 57.31% in 2012 and 47.82% and 52.38% in 2013, respectively, compared to those under AT. In terms of the dry matter contribution to the grain of the stem and sheath, the difference between the AT and MT varied with the varieties due to the different genotypes. In 2012, the dry matter contribution of the stem and sheath for Shengdao2572 and Jindao263 under MT significantly decreased, while there was no significant difference between AT and MT for Daliang202 and Huaidao11. In 2013 under MT, this value significantly decreased for most of the varieties excluding Daliang202 (Table 4).

Table 4. Above-ground dry matter remobilization characteristics of rice under AT and MT in 2012 and 2013.

Leaf Stem and sheath
Year Cultivar Cultivation pattern Dry matter remobilization amount (g) Dry matter remobilization efficiency (%) Dry matter contribution to grain yield (%) Dry matter remobilization amount (g) Dry matter remobilization efficiency (%) Dry matter contribution to grain yield (%)
2012 Shengdao2572 AT 0.342a 44.0a 17.9a 0.0578a 6.251a 3.022a
MT 0.137b 28.5b 12.2b 0.0166b 2.336b 1.489b
Daliang202 AT 0.360a 40.8a 18.0a 0.1248a 11.715a 6.229a
MT 0.173b 25.0b 14.0b 0.0734b 8.343b 5.973a
Huaidao11 AT 0.322a 36.4a 13.9a 0.1240a 11.403a 5.347a
MT 0.110b 16.8b 7.1b 0.0717b 7.260b 4.558a
Jindao263 AT 0.320a 39.7a 13.0a 0.1317a 10.560a 5.274a
MT 0.176b 28.5b 11.0b 0.0318b 2.895b 1.980b
2013 Shengdao2572 AT 0.291a 43.1a 17.4a 0.049a 5.9a 2.9a
MT 0.094b 26.4b 10.4b 0.011b 2.1b 1.2b
Daliang202 AT 0.444a 56.1a 24.7a 0.113a 12.1a 6.4a
MT 0.259b 37.5b 21.1b 0.081b 9.2b 6.6a
Huaidao11 AT 0.246a 36.3a 11.5a 0.107a 10.8a 5.0a
MT 0.081b 13.2b 4.7b 0.036b 3.9b 2.1b
Jindao263 AT 0.246a 36.1a 11.3a 0.066a 6.9a 3.0a
MT 0.135b 18.6b 8.3b 0.015b 1.8b 0.9b

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

The change trends of the ratios of the leaf, stem and sheath, and panicle dry matter weight to the total weight at the heading stage under MT were different from those under AT across the two study years. The ratio of the leaf dry matter weight to the total weight under MT markedly decreased by 15.13% except for cultivar Jindao263 in 2012, while it decreased by 6.25% except for in Huaidao11 and Jindao263 in 2013. In 2012, the ratio of the dry matter weight of the stem and sheath to the total weight significantly decreased in Shengdao2572 and Huaidao11 under MT but not in the other varieties. However, this difference was insignificant between AT and MT for most of the cultivars in 2013. In 2012, the ratio of the panicle dry matter weight to the total weight under MT significantly increased in Daliang202 and Huaidao11 but showed little change in the other two cultivars. In 2013, this ratio significantly increased in Daliang202, Huaidao11 and Jindao263, while the opposite occurred in Shengdao2572. Compared to AT, the ratios of the leaf, and stem and sheath dry matter weight to the total weight in every variety at maturity under MT significantly increased by 19.56% and 17.52% in 2012 and 27.63% and 10.31% in 2013, respectively. In contrast, the ratio of panicle dry matter weight to the total weight under MT markedly decreased by 15.03% and 12.93% in 2012 and 2013, respectively (Table 5).

Table 5. Above-ground dry matter distribution of rice under AT and MT in 2012 and 2013.

Ratio of the dry weight of the leaf to the total dry weight (%) Ratio of the dry weight of the stem and sheath to the total dry weight (%) Ratio of dry weight of the panicle to the total dry weight (%)
Year Cultivar Cultivation pattern heading stage Maturity stage heading stage Maturity stage Heading stage Maturity stage
2012 Shengdao2572 AT 37.1a 13.5b 44.1b 26.9b 18.8a 59.5a
MT 33.0b 16.0a 49.1a 32.4a 17.9a 51.6b
Daliang202 AT 40.2a 15.1b 48.4a 27.0b 11.4b 57.9a
MT 31.7b 20.3a 46.5b 31.5a 16.9a 48.2b
Huaidao11 AT 37.7a 14.6b 46.5b 25.0b 15.8b 60.4a
MT 32.9b 18.1a 49.2a 30.2a 17.9a 51.7b
Jindao263 AT 33.6a 11.9b 51.4a 27.0b 14.9a 61.1a
MT 33.3a 14.1a 50.4a 34.3a 16.3a 51.5b
2013 Shengdao2572 AT 36.1a 13.5b 44.4b 27.5b 19.5a 59.0a
MT 33.9b 15.6a 48.4a 29.9a 17.3b 54.5b
Daliang202 AT 39.1a 11.8b 46.4a 27.8b 14.5b 60.4a
MT 36.6b 17.6a 46.2a 32.3a 17.3a 50.1b
Huaidao11 AT 34.6a 12.5b 50.3a 25.5b 15.1b 62.0a
MT 33.1a 17.0a 49.7a 28.2a 17.2a 54.8b
Jindao263 AT 35.9a 12.5b 49.9a 25.3b 14.2b 62.2a
MT 34.5a 19.3a 49.0a 27.9a 16.5a 52.7b

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

Population photosynthetic ability and growth rate for rice from old seedlings under MT

Compared to that under AT, leaf area index at the jointing, booting, heading and maturity stages under MT significantly decreased by 48.05%, 35.36%, 37.11% and 58.01% in 2012 and 48.55%, 35.76%, 32.58% and 57.54% in 2013, respectively; The reduction rate of the leaf area from heading to maturity under MT significantly decreased by 10.35% in 2012 and 7.92% in 2013, respectively. In 2012, the photosynthetic potential and crop growth rate under MT significantly decreased by 43.31% and 25.49% from jointing to booting, 51.81% and 34.51% from booting to heading, and 47.06% and 46.66% from heading to maturity. In 2013, these values were 40.07% and 39.78% from jointing to booting, 37.52% and 35.92% from booting to heading, and 42.18% and 37.17% from heading to maturity, respectively (Table 6).

Table 6. Leaf area index, reduction rate of the leaf area, photosynthetic potential and crop growth rate of rice under AT and MT in 2012 and 2013.

Leaf area index Reduction rate of leaf area (LAI d-1) Photosynthetic potential (m2 d) Crop growth rate (g m-2 d-1)
Year Cultivar Cultivation pattern Jointing Booting Heading Maturity from heading to maturity From jointing to booting From booting to heading From heading to maturity From jointing to booting From booting to heading From heading to maturity
2012 Shengdao2572 AT 4.015a 6.529a 6.389a 3.933a 0.052b 57.991a 64.593a 242.604a 16.419a 24.252a 8.936a
MT 2.204b 4.152b 4.140b 1.711b 0.057a 34.960b 33.168b 125.806b 8.819b 9.306b 4.899b
Daliang202 AT 4.558a 6.160a 7.203a 4.392a 0.057b 69.669a 66.815a 284.082a 9.812a 28.405a 8.841a
MT 2.338b 4.251b 4.749b 2.032b 0.062a 46.124b 31.502b 149.199b 4.809b 23.580b 6.276b
Huaidao11 AT 5.137a 6.661a 7.250a 5.221a 0.046b 82.582a 76.506a 274.341a 24.623a 32.133a 11.897a
MT 2.425b 4.052b 4.144b 1.798b 0.051a 42.100b 32.784b 136.672b 22.941b 15.155b 5.806b
Jindao263 AT 5.632a 7.949a 7.775a 5.616a 0.048b 101.857a 78.618a 301.296a 23.484a 43.144a 14.270a
MT 3.081b 5.190b 4.965b 2.506b 0.054a 53.762b 40.621b 171.844b 18.824b 35.746b 6.458b
2013 Shengdao2572 AT 3.725a 5.849a 5.579a 3.333a 0.047b 38.296a 51.426a 213.890a 22.913a 16.740a 7.791a
MT 1.977b 3.836b 3.856b 1.396b 0.052a 29.068b 34.614b 123.417b 6.132b 3.321b 4.421b
Daliang202 AT 4.815a 7.559a 7.226a 4.765a 0.052b 61.869a 59.299a 288.741a 20.742a 32.437a 8.238a
MT 1.962b 4.778b 4.531b 1.847b 0.056a 37.069b 41.891b 153.073b 10.370b 23.404b 5.553b
Huaidao11 AT 5.075a 6.953a 6.282a 3.826a 0.051b 66.156a 66.174a 242.593a 22.241a 26.647a 8.599a
MT 3.108b 4.127b 4.168b 1.522b 0.054a 32.556b 29.030b 139.392b 16.381b 22.965b 5.344b
Jindao263 AT 5.239a 7.195a 6.718a 4.193a 0.052b 74.605a 62.610a 267.321a 42.093a 11.675a 7.452a
MT 2.654b 4.961b 4.843b 2.078b 0.056a 45.691b 44.119b 169.566b 32.153b 6.381b 4.837b

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

Performance of the agronomic traits and grain yield for rice from old seedlings under MT

As shown in Fig 3, a significant difference in the plant height at maturity between AT and MT was found. Except for cultivar Huaidao11, the plant height for the other cultivars under MT was 8.79% less than that under AT in 2012. Similarly, the plant height significantly decreased by 9.85% in 2013.

Fig 3. Rice plant height at maturity under AT and MT in 2012 and 2013.

Fig 3

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

Compared to AT, there was a significant decrease in the panicle length under MT in 2013; however, the difference between both of the treatments was not significant except for that in cultivar Shengdao2572 in 2012. The number of primary branches per panicle markedly decreased by 11.12% and 15.65% under MT, except for Huaidao11 in 2012 and Daliang202 and Huaidao11 in 2013. Under MT, the number of the secondary branches per panicle significantly decreased by 36.10% and 27.56%, excluding Shengdao2572 and Huaidao11 in 2012 and Huaidao11 in 2013. In 2012, the seed-setting rate markedly decreased except for that in Daliang202 under MT compared to that under AT. Nevertheless, no significant difference between the two treatments was observed except for in Shengdao2572 in 2013. There was a difference in the 1000-grain weight among the cultivars and study years between AT and MT. In 2012, the 1000-grain weight of Shengdao2572 under MT significantly decreased, while the opposite occurred for Jindao263, with little alteration being found in Daliang202 and Huaidao11. In 2013, the 1000-grain weight significantly increased under MT in Shengdao2572 and Jindao263 but not in the other two cultivars. The effective panicles per mm2, total grains per panicle (except Huaidao11) and grain yield significantly decreased by 28.97%, 18.34% and 31.26%, respectively, in both of the study years (Table 7).

Table 7. Agronomic traits of the panicle, grain yield and yield components in rice under AT and MT in 2012 and 2013.

Year Cultivar Cultivation Pattern Panicle length (cm) No.of the primary branch per panicle No. of the secondary branch per panicle No. of effective panicles m-2 No. of total grainspanicle-1 Seed-setting rate (%) 1000-grain weight (g) Grain yield(kg m-2)
2012 Shengdao2572 AT 16.45a 10.73a 19.57a 380.80a 98.13a 0.89a 26.72a 0.66a
MT 14.64b 9.70b 17.17a 265.72b 78.45b 0.77b 25.73b 0.46b
Daliang202 AT 16.02a 12.30a 26.82a 438.76a 120.96a 0.93a 25.97a 0.97a
MT 15.89a 10.93b 17.70b 358.68b 95.45b 0.90a 26.50a 0.62b
Huaidao11 AT 16.26a 11.51a 17.36a 396.76a 109.94a 0.91a 27.20a 1.10a
MT 16.40a 11.50a 17.90a 242.76b 106.79a 0.77b 27.37a 0.60b
Jindao263 AT 16.09a 12.31a 20.71a 620.76a 110.82a 0.95a 25.55b 1.17a
MT 16.40a 10.78b 12.67b 294.00b 99.99b 0.84b 27.27a 0.80b
2013 Shengdao2572 AT 18.23a 11.67a 22.39a 346.92a 131.11a 0.78a 23.64b 0.80a
MT 16.65b 10.55b 19.40b 263.76b 109.00b 0.73b 27.12a 0.43b
Daliang202 AT 17.14a 11.67a 25.90a 399.00a 118.27a 0.90a 25.69a 1.25a
MT 14.75b 11.10a 18.91b 317.24b 105.86b 0.93a 25.68a 0.93b
Huaidao11 AT 17.97a 11.43a 21.00a 406.00a 120.61a 0.91a 28.29a 0.94a
MT 16.60b 10.72a 21.20a 284.76b 123.78a 0.90a 27.46b 0.82b
Jindao263 AT 17.85a 12.93a 23.37a 426.16a 145.28a 0.89a 26.50b 1.14a
MT 15.57b 10.20b 13.60b 399.00b 102.93b 0.87a 27.15a 0.86b

Values of AT and MT for the same cultivar followed by different letters represent significant difference at p≤0.05 level.

Relationships between the grain yield and dry matter production, remobilization and distribution characteristics

There existed a significant and positive correlation between the dry matter weight per tiller at the jointing-maturity stage, the population photosynthetic potential, the crop growth rate, the dry matter remobilization efficiency at the heading-maturity stage, the panicle dry matter distribution ratio at maturity and the grain yield. Furthermore, there was a significant and negative correlation between the leaf dry matter distribution ratio and the stem and sheath dry matter distribution ratio and grain yield. These relationships were well simulated by linear equations, which were presented in both of the study years (Fig 4).

Fig 4. Relationship between grain yield and the dry matter per tiller, the photosynthetic potential of the population, the crop growth rate, the dry matter remobilization efficiency and the above-ground dry matter distribution ratio at maturity.

Fig 4

Discussion

Delayed rice growth stages under MT compared to under AT were previously reported by Huo [19], who demonstrated that the rice-jointing, heading and maturity stages were delayed by 5–8 d, 2–3 d, and 1–2 d, respectively, when the rice seedlings were transplanted at the optimum seedling age (approximately 20 d). This study demonstrates that, compared to those under AT, rice jointing, booting, heading and maturity stages were remarkably delayed by 8–12 d, 6–13 d, 5–9 d and 5–6 d under MT in 2012, respectively, while they only had a marginal change in magnitude with a delay of 2–7 d under MT in 2013. The difference between both of the study years can be explained as higher air temperature during rice growth in 2013, especially from the booting stage to the heading stage (Table 1 and Fig 1). The delayed rice growth under MT may be attributable to the deleterious effects on the roots of mechanical transplantation. In general, it takes rice plants approximately 10–14 d to recover from injury. The age of the seedlings at transplantation significantly affects the number of productive tillers [20]. When rice seedlings are transplanted at the right time in terms of age, tillering and growth proceed normally [21]. The occurrence of tillering for rice with 20-d-old seedlings under MT commenced from the third leaf position after the seedlings were moved into a paddy field, and the peak number of tillers was much greater under MT than that under AT [22]. In contrast to those under AT, the rice tillers under MT occurred 7–10 d later but increased more rapidly once normal growth resumed after being transplanted [22]. In the experiment, the occurrence of rice tiller under MT began almost from the 12th day after being transplanted. It is noteworthy that the maximum and increasing rates of rice tillers under MT were markedly smaller than were those under AT, which is inconsistent with the report of Li [22]. This contradiction may be ascribed to the intensive competition among individual seedlings under the limited growth conditions of the seed-nursery bed under MT, which lasts longer for old seedlings than that for normal seedlings with optimum seedling age [21]. As a result, the tiller occurrence in rice from old seedlings under MT was severely hampered by the delay in transplantation, i.e., the later the rice seedlings were transplanted by machine, the more severely was hindered the rice tiller growth.

The above-ground dry matter weights per tiller and population are pivotal parameters reflecting the growth potential of individuals and populations of rice. The population dry matter weight is determined by the tiller number and dry matter accumulative amount per tiller. Therefore, realizing the growth potential of individuals is beneficial to the improvement of the ability to produce dry matter for the rice population. A previous study reported that the dry matter weight per tiller for mechanically transplanted 20-d-old rice seedlings was 13.15%, 7.73%, 8.77% and 10.84% lower than that of rice that was transplanted by hand at the jointing, booting, heading and maturity stages, respectively [24]. In terms of the dry matter accumulative amount of the population, significant differences between MT and AT were detected, with the former being 6.95% and 11.00% lower than the latter from jointing to heading and from heading to maturity [23]. In the present study, the dry matter weight of the leaf or the stem and sheath from jointing to maturity as well as that of the panicle from heading to maturity in rice (seedling age 42 d) under MT significantly declined compared to that under AT. In addition, a similar tendency also occurred in the dry matter accumulative amount of the rice population from jointing to booting, from booting to heading and from heading to maturity (Tables 1 and 2), which seems to be similar to the reported results [23]. Nevertheless, the differences between AT and MT expanded in our study compared to those in the above-mentioned report, suggesting that the growth and development of rice plants under MT was evidently retarded with the delayed transplantation.

In general, rice grain-filling matter consists of nonstructural carbohydrates that are stored in the stem, sheath and leaves pre-anthesis and of photosynthetic products created mainly by leaves post-anthesis, with the former accounting for nearly 30% [23]. Regarding the rice plants that were transplanted by machine at a 20-d seedling age, the dry matter remobilization amount and efficiency and the contribution rate to the leaf yield decreased by 21.41%, 13.16% and 14.35%, respectively, whereas these values increased by 11.06%, 20.48% and 21.16% for the stem and sheath compared to rice plants that were transplanted by hand [23]. According to the data of the present study, under MT, the above-mentioned parameters (except for the contribution rate to the yield for the stem and sheath) were significantly lower than were those under AT (Table 4). These results indicate that the supply of photosynthetic assimilates from source (leaf + stem and sheath) to sink (grain) is lowered for rice plants from old seedlings under MT, which may be linked to decreased sink capacity (Table 7). The photosynthetic potential is a key index representing the photosynthesis productivity of the crop population, which depends on both the leaf index and the duration of leaf photosynthesis [25]. The growth rate of a crop population, a parameter embodying the dry matter production per day, describes the efficiency of photosynthesis productivity [26]. Our results demonstrate that, in contrast to those under AT, the rice photosynthetic potential and the population growth rate at the jointing-heading and heading-maturity stages significantly decreased under MT (Table 6). The differences between AT and MT are greater than those that were reported by Li [23], who found that under MT, the rice (20-d seedling age) photosynthetic potential at every growth stage was 11.19% lower than that under AT, while the population growth rate was slightly lower than that under AT from the jointing to the heading stage (with a significant decrease only from heading to maturity). One possible explanation for the difference in these agronomic traits between AT and MT is the shortage of vegetative growth due to the inhibited growth in the nursery bed and the postponed recovery from mechanical transplantation injury for mechanically transplanted rice, which was also evidenced by the decreased plant height and tiller occurrence (Figs 2 and 3). Furthermore, if the rice plants are mechanically transplanted after the appropriate seedling age, the vegetative growth duration of rice is further shortened due mainly to the longer growth inhibition in the nursery bed, which demonstrates that the development of the tiller, the photosynthetic production and the assimilate remobilization efficiency in rice from old seedlings are severely impaired. These speculations demonstrate that the above-ground characteristics of rice plants after transplanting varied with seedling age.

Seedling age is an important element affecting the number of filled grains per panicle, the panicle length, the 1000-grain weight and the grain yield in rice, as reported by Ginigaddara [20]. The timely transplantation and appropriate seedling age are essential for producing a higher grain yield, while the seedling age depends on many factors, including crop rotation, climate condition, etc., and might be different in different production systems [27,28,29,30]. Li [23] reported that, when rice plants were mechanically transplanted at a 20-d seedling age, the grain yield markedly decreased by 4.66% compared to that under AT. However, in the present study, the rice (42-d seedling age) grain yield under MT was 31.26% lower than that of AT (Table 7). These results indicate that delaying transplanting has a significant negative influence on the rice grain yield under MT. In addition, our results indicate that the number of effective panicles per m2 under MT significantly decreased compared to that under AT, and a similar trend was observed in the number of primary and secondary branches and the total grains per panicle (except for in the cultivar Huaidao11) (Table 7). The significant decrease in the number of effective panicles is largely attributed to poorer tiller occurrence under MT (Fig 2). Therefore, we argue that the significant reduction in the number of effective panicles and filled grains per panicle is the direct cause of the decreased grain yield under MT. This study reported that high-yielding rice was characterized by great population dry matter weight and photosynthetic potential and a slowed leaf-area reduction rate. After the heading stage, transferring more photosynthetic product from being stored in the leaf, stem and sheath to the panicle allows rice to obtain a high grain yield; in other words, a higher dry matter distribution ratio of the panicle is instrumental in increasing the grain yield. Our results indicate that dry matter weight per tiller at the jointing-maturity stage, the population photosynthetic potential, the crop growth rate, the dry matter remobilization efficiency and the panicle dry matter distribution ratio are significantly and positively correlated with grain yield, while the leaf dry matter distribution ratio and the stem and sheath dry matter distribution ratio are significantly and negatively related with the grain yield (Fig 4). Therefore, we speculate that the decreased photosynthetic potential, population growth rate, dry matter accumulative amount, dry matter distribution ratio of the panicle and accelerated reduction of the leaf area are important physiological factors in the decreased rice grain yield under MT. Interestingly, compared to that under AT, the 1000-grain weight of cultivar Jindao263 under MT significantly increased irrespective of the impaired photosynthetic productivity and assimilate supply. Presumably, the sink capacity (effective panicles and filled grains) decreases more than does the source supply (photosynthetic productivity); in other words, the source-sink ratio increases, thereby leading to an increased grain weight in Jindao263.

Compared with AT, the tiller numbers, dry matter weight, dry matter accumulation, LAI, growth rate, photosynthetic potential, dry matter remobilization efficiency of leaf under MT were significantly decreased under MT. Those agronomic traits above are attributed to grain yield reduced under MT. Taken together, delaying transplanting severely restricted rice growth and grain yield under MT in the Huanghuai region of China. Consequently, it is crucial to adopt effective measurements to mitigate the phenomenon. Firstly, altering the settings of the transplantation machine to lessen damage during transplantation may be an alternative in shortening resuming growth period and expediting rice seedling tillering. Secondly, lowering sowing density could decrease the competition among plant individuals in nursery bed and improve rice seedling quality. Improving the settings of the transplantation machine could match the lowered sowing density. Thirdly, screening varieties with relative insensitivity to delaying transplanting and consistent performance in yield traits may be also an effective channel.

Conclusions

This investigation indicates that using old rice seedlings would result in postponed growth stages, poor tiller occurrence, limited dry matter accumulation and lower population growth rate, photosynthetic potential, and grain yield under MT. Compared to those under AT, the tiller number, population photosynthetic productivity, and dry matter remobilization efficiency significantly decreased under MT due primarily to the obviously short vegetative growth duration, which is the main physiological reason for the decreased grain yield. The significant decreases in the filled grains per panicle and in the number of effective panicles in particular directly led to a decreased rice grain yield, especially that of effective panicles.

Data Availability

All relevant data are within the paper.

Funding Statement

The authors received no specific funding for this work.

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Associated Data

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Data Availability Statement

All relevant data are within the paper.


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