Timelines of [21]rice breeding activities 2006–2012.
\r\n\tIt is a relatively simple process and a standard tool in any industry. Because of the versatility of the titration techniques, nearly all aspects of society depend on various forms of titration to analyze key chemical compounds.
\r\n\tThe aims of this book is to provide the reader with an up-to-date coverage of experimental and theoretical aspects related to titration techniques used in environmental, pharmaceutical, biomedical and food sciences.
Rice is an important food crop that is consumed mostly outside its major production areas in Uganda, with over 90% of production marketed to urban areas and major institutions within the country. This aspect makes rice to have a long value chain engaging several players. Rice is cultivated under rain‐fed upland conditions, partly rain‐fed lowland conditions and irrigated conditions in Uganda, taking advantage of diverse ecosystems in Uganda [1]. Since the introduction of rice in 1904, Uganda had production under different agro‐ecological conditions covering rain‐fed upland conditions, partly rain‐fed lowland conditions and irrigated conditions. Various production challenges are faced in these production areas.
\nNew technologies are valuable for use in developing the new rice varieties globally including Africa. However, new tools can be most helpful if the existing varieties and candidate lines are properly characterized and documented. The purpose of this paper is to trace upland rice breeding efforts in Uganda and present for alignment, learning, and application in the new technology in their rice breeding programs with focus on breeding for drought tolerance and other stresses. This is critical considering that there is limited research on development of upland rice varieties suitable for production under mild drought conditions in the East African region and other similar Agro ecologies. There is also increasing expansion of rice production from traditional irrigated production areas to rain‐fed environments, where drought problem is an inherent challenge. Indeed, drought emerged as a critical rice production constraint in East Africa [1, 2], particularly in Uganda [1], as promotion of upland rice was growing in the country.
\nMany upland rice varieties, earlier introduced in the country, were late maturing and did not have preferred cooking qualities. Later, more introduced lines were evaluated and released. These varieties had been generated through interspecific crossing involving Oryza glaberrima and Oryza sativa. These new genotypes were called the ‘New Rice for Africa’ (NERICA). They were resistant to major biological constraints but showed differential sensitivity to drought stress and new diseases, especially brown spot disease and narrow leaf spot disease. Besides, these varieties had nonaromatic characteristic which are the major concerns of the Uganda farmers. These factors made upland rice farmers to realize low yield mainly due to frequent drought stress. In addition, extensive use of irrigated rice come along with other limitations, namely need for environmental impact assessment and conflict on cultural values for use of the wetlands for farming. Subsequently, upland rice breeding involving adapted varieties led to new rice varieties with high resistance to biotic stresses and preferred agronomic traits [3]. However, abiotic stresses, especially drought stress remained a major constraint. Indeed, breeding for drought tolerance resistance in rice is challenging because the trait is quantitative and involves polygenes with low heritability. Modified pedigree breeding approaches were used in this breeding. In this paper, we review a trend of improvement of upland rice in Uganda covering three aspects: (1) screening of introductions for drought tolerance, (2) mode of gene action for drought tolerance, (3) evaluation of segregating lines, (4) Evaluation of promising lines, (5) variety release and status of deployment of the new generations of rice varieties in Uganda and within the African region. Detailed timelines of the activities are presented in Table 1.
2006B | 2007A | 2007A | 2007B | 2008A | 2008B | 2009A | 2009B | 2010A | 2010B | 2011A | 2011B | 2012A | 2012B | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
New lines bred in Uganda | ||||||||||||||
Selection of parental lines | ||||||||||||||
Making crosses and generating F2 (genetic studies) | ||||||||||||||
Evaluation of F2 and harvesting F3 | ||||||||||||||
Evaluating F3 under rain‐fed lowland and upland conditions (660) | ||||||||||||||
Evaluation F4 upland conditions | ||||||||||||||
Evaluation F5 in Advanced Trials | ||||||||||||||
Fixed lines introduced | ||||||||||||||
Evaluation F6 in NPT (12 lines) | ||||||||||||||
Evaluation F6 in NPT (10 lines) | ||||||||||||||
Testing for grain traits | ||||||||||||||
Organoleptic testing of the new varieties | ||||||||||||||
PVS selection of varieties |
Timelines of [21]rice breeding activities 2006–2012.
A total of 191 rice introductions from major rice breeding centers were evaluated. Of the 191 materials, 77 were O. sativa indica comprising 45 from African Rice Centre (ARC), 15 lines from International Rice Research Institute (IRRI), 13 from Mali, three from Uganda, and one from China. Among the introductions, there were three O. glaberrima accessions. The remaining 111 were interspecific lines developed from O. sativa × O. glaberrima crosses, comprising 18 from ARC and 93 the International Center for Tropical Agriculture (CIAT), Colombia coded as the CT series. However, among the interspecific samples, two genotypes namely WAB 880‐1‐27‐9‐2‐P1‐HB and WAB 450‐24‐2‐3‐P‐38‐1‐HB were duplicates from different repeated introductions from IRRI and WARDA‐Africa Rice Center. The 93 interspecific lines from CIAT were BC4F1s developed from crossing CAIAPO, a tropical O. sativa japonica from Colombia with RAM 24 (O. glaberrima).
\nThis experiment was conducted at National Crop Resources Institute (NaCRRI), at Namulonge in central Uganda, at 00°32’ N latitude and 32°53’ E longitudes with altitude of 1150 m above sea level during dry season. The soils of the place are clay loam. The period December to March is characteristically the long dry season, but mean long term annual rainfall is 1270 mm.
\nIn order to assess drought stress during reproductive growth stage, drought stress was imposed by terminating irrigation, when about 50% of the population had reached a point where interauricular distance between the flag leaf and penultimate leaf was zero [4]. It is the period when it is about 10 days before anthesis. It is the time when the penultimate leaves were fully expanded. Rainfall during the trial period was recorded. Irrigation was 14 days later, when 30% of the available water had been lost from the soil at 20‐cm depth. The available soil moisture was taken using the ECHO soil moisture tester (Decagon Devices, Inc., Pullman, Washington, USA). All the grains from each panicle were hand threshed and dried. The filled and unfilled grains were then separated using floatation methods.
This study investigated the nature of inheritance of drought tolerance in crosses between interspecific and intraspecific rice genotypes using secondary traits. Two separate experiments were conducted, using O. sativa and fixed interspecific lines derived from O. glaberrima and O. sativa crosses.
\nExperiment 1: Genetic studies on drought tolerance traits
\nThe aim of the first experiment was to investigate the inheritance of drought tolerance at reproductive growth stage. Eighteen crosses were generated from two sets of 3 × 3 parents using the North Carolina mating design II (NCD II). All the 18 F2 and the 12 parents were evaluated in a 2 × 15 alpha lattice design with two replicates under a rain‐out shelter and nonstress conditions in the field.
\nThirty genotypes comprising 18 F2 progenies from sets, A and B, along with the 12 parents were used in this experiment (Table 2). The 30 entries were established in a rain‐out shelter at National Crops Resources Research Institute (NaCRRI), Namulonge. The rain‐out shelter was constructed using translucent sheets for the roof and wire mesh on the sides of the structure to prevent rain water and to allow free air circulation, respectively. In the rain‐out shelter, standard troughs that are 1m wide, 8m long, and 1.5m deep were filled with soil for fallow field from Namulonge. Four troughs were made and filled with the soil, referred to as strips. The seeds were planted in a 2 × 15 alpha lattice design. Two strips represented a replicate. The 12 parental genotypes were planted in three rows planted across the 1 m width strips, while the F2 populations were planted in six rows. The plant to plant spacing was 15 cm making plant population to be 36 for the parental lines and 72 for the F2 lines.
Experiment | Crossing set | Genotype no | Breeding line | Type | Parent type |
---|---|---|---|---|---|
1 | SET A | 18 | CT 16334(2)‐CA‐2‐M | Interspecific | Male |
105 | WAB 365‐B‐1H1‐HB | O. sativa | Male | ||
134 | NERICA 9 | Interspecific | Male | ||
138 | NERICA 8 | Interspecific | Female | ||
193 | NERICA 13 | Interspecific | Female | ||
196 | IRAT 325 | O. sativa | Female | ||
SET B | 2 | CT 16346‐CA‐20‐M | Interspecific | Male | |
9 | CT 16350‐ CA‐5‐M | Interspecific | Male | ||
12 | CT 16344‐CA‐9‐M | Interspecific | Male | ||
96 | Bonanca | O. sativa | Female | ||
121 | WITA 2 | O. sativa | Female | ||
129 | CK 73 | O. sativa | Female | ||
2 | SET C | 18 | CT 16334 (2)‐CA‐2‐M | Interspecific | Female |
138 | WAB 450‐1‐BL1‐136‐HB | Interspecific | Male |
Rice genotypes used for generating sets of F1 for drought tolerance.
A second set of the 30 entries were planted in the field under optimal conditions. These conditions involved irrigating the field at 20 mm per week, during the period when there was no rain. In both trials, a 2 × 15 alpha lattice design planted in two replicates was used. Two seeds from each generation were drilled at a depth of 3 cm at spacing of 20 × 20 cm in each plot. In order to reduce border effects, 20 cm was left between plots. The 12 parental genotypes were planted in 5‐row and 3‐column plots, while the F2 populations were planted in 5‐row and 6‐column plots. Overall, there were 15 plants per replicate of the parents and 30 plants per replicate of each F2 genotype, thus the total number of plants were 30 and 60 for the parents and F2, respectively. The plants were thinned to one plant per hill. Standard cultural practices including hand planting and hand weeding were followed. The crops were fertilized with 25 kg N ha-1 at 20–25 days after transplanting (DAT) and the same rate at 40–45 DAT to enhance plant vigor.
\nDrought stress was imposed by terminating irrigation, when about 50% of the populations had attained an interauricular distance between the flag leaf and penultimate leaf of zero, that is the period about 10 days before anthesis [4, 5]. This method of identifying the stage of imposing drought was applied both in the field and in the rain‐out shelter. In general, this is the time when the penultimate leaves were fully expanded. Rainfall during the trial period was recorded.
\nIn the field experiment, irrigation was applied using sprinkler irrigation. The field was irrigated, every three days before imposing drought stress. On the day the irrigation was terminated, the field was irrigated to field capacity in the evening between 5:00 and 6:00 pm, which was resumed 14 days after its termination using sprinkler irrigation. The duration of drought stress was determined by testing the level of soil moisture daily, using the ECHO soil moisture tester (Decagon Devices, Inc Pullman, Washington USA). On the day, when 30% of the available water had been lost from the soil at 20‐cm depth, irrigation was resumed. In the rain‐out shelter, water was applied using hand irrigation cans but water was calculated for each strip at 140 L per week, which is equivalent to 20 mm per week.
\nThe number of filled grains was counted per panicle at grain maturity period. Two panicles from each plant were randomly collected and record of number of filled grains was determined using floatation method described by these authors [6].
\nExperiment 2: Generation means analysis (GMA) for filled grains in rice
\nIn the second experiment, the magnitude and direction of gene action for drought tolerances at reproductive stage was determined in five populations P1, P2, F1, F2, and F3 generated from a drought tolerant × susceptible cross using generation mean analysis (GMA). They are in set C (Table 2). The materials were planted in the dry season and drought was imposed by terminating at the stage of panicle initiation.
\nIn this experiment, all the five populations generated from crossing; parents P1 and P2, their F1, F2 and F3 genotypes were planted following a randomized complete block design (RCBD) with two replicates. Two seeds from each generation were drilled at a depth of 3 cm at spacing of 20 × 20 cm in each experimental unit (plot) in the field at NaCRRI. The generations P1, P2, and F1 were planted in 5‐row and 3‐column plots, while F2 and F3 were planted in 5‐row and 6‐column plots. Overall, there were 15 plants per replicate of the parents, 30 plants per replicate of each F2 genotype, thus the total numbers of plants were 30 and 60 for the parents, F2, respectively. The cultural practice in experiment 1 was followed, and drought stress was imposed following procedures in experiment 1.
\nData was analyzed in three parts, namely analyses of variance, residual maximum likelihood (REML), regression, and generation means. The analysis of variance was performed for different traits associated with drought tolerance in the two sets of populations, A and B, pooling for both stress and nonstress environments. Using REML, the separate sets were analyzed for each trait. The analyses of the variance components of genotypes were further partitioned into variations, due to parents and crosses.
\nGeneral analyses of variance were performed for filled grains, grains per panicle, leaf area, plant height, tiller number, and panicle number of all hybrids including checks. Genetic analyses for the six parameters of experimental hybrids were then performed in GenStat [7] as a fixed effects model across two locations [8] as follows:
\nGeneration mean analysis of the genotypes CT 16334 (2)‐CA‐2‐M crossed with WAB 450‐1‐BL1‐136‐HB was used to determine additive, dominant, and epistatic effects following the model [9]. The various generations did not have equal variances; therefore, weighted inverse of the variances was used in subsequent analysis according to these authors in Ref. [10]. Regression analysis procedures were used to find the best fit model. It is a graphical method used to compare the additive model with additive‐dominance models. Any effect that was not significant at 5% level was excluded from the model. The parameters were fitted using weighted mean squares as described by Ref. [11].
\nA scaling test was conducted using linear combinations of various means according to Refs. [9–12] to detect the presence of nonallelic interactions that are known to bias estimates of additive and dominance components in the populations when present. However, in this case, where F3 populations are used instead of backcross populations, the additive effects estimate is for both additive effects and additive × additive interaction effects. Similarly, the dominance effect combined both dominance effects and dominance × dominance interaction effects as a single estimate [12]. This is not a major drawback considering that most breeding work exploits additive effects and dominance effects. Standard errors of generation means were computed by performing nested analysis of variance following methods used in Ref. [13].\x3c!-- References were renumbered because they are out of sequence. Please check if changes have been carried out accurately.
In order to verify the number of genes involved in the transmission of traits associated with drought tolerance, Castle‐Wrights formulae described in Ref. [14] was used.
Preliminary yield trials were conducted on station with objective of varietal screening, evaluation, and seed increase. These were F3 selections from the previous experiment. Overall 660 genotypes were selected from the F3 generation based on field performance. All the seed from each of the 660 hills of F3 genotypes were divided into three sets. One set was remnant; a second set was planted in the rain‐fed low‐land environment, while the third set was planted under rain‐fed upland conditions, all on station within Namulonge. The planting was in November, 2010. All the seed from each hill was planted to 5‐m long rows and evaluated.
\nThe second set was planted under rain‐fed low‐land with ample moisture throughout the growth period of the lines. The evaluation focused on maturity period, tillering capacity, presence of foliar diseases, and physical grain characteristics. Lines that had longer maturity period than the variety NERICA‐4, number of reproductive tillers less than 5 per hill, presence of foliar diseases, and grain discoloration were eliminated. Besides, infection by common pathogen namely rice blast, bacterial leaf blight, grain discoloration, and sheath rot were used to eliminate lines. The team that evaluated the materials comprised of scientists, farmers, and rice field workers.
\nThe third set comprising all the 660 lines was planted under rain‐fed upland conditions. Selection was made as previously stated for rain‐fed low‐land conditions. Unlike selection under rain‐fed lowland conditions where a minimum of seven productive tillers was considered acceptable, in this production environment, five productive tillers was considered the minimum.
Set 1: Evaluation of 84 rain‐fed lowland rice lines: a total 84 lines of F4 segregating populations were selected from the 660 F3 lines genotypes and planted in five sites namely Namulonge, Kigumba, Kibaale, Lira, and Doho. Each entry was planted in a 3 × 18 alpha lattice design at spacing of 20 × 20 cm planted in rows five plots each 5‐m long. The evaluation had three main objectives. The first objective was to test the new genotypes under varying stress conditions. The major biotic and abiotic stresses targeted were drought stress, rice blast, RYMV, BLB and Leaf Streak, narrow leaf spot and brown leaf spot. The locations selected were major rice growing areas that had had the production constraints. The second objective was to assess yield of the whole set at Namulonge site. The third objective was to identify farmer preferred varieties using participatory variety selection method.
On‐farm trials were conducted through participatory and multilocational testing of selected upland varieties. Selection of sites for participatory and multilocational testing considered the following: (i) key representative ecological zones, (ii) participation of stakeholders, and (iii) availability of resources to effectively conduct the exercise. Seed companies were invited and a proposed method of allowing most seed stakeholders to participate in variety evaluation was adhered to. Twenty lines that were tested in 2011B in two locations namely Namulonge and Kibaale. Subsequently, 12 lines were tested in 2012A and finally 8–10 in 2012 B.
\nIn order to identify suitable upland rice varieties, the best rice lines from preliminary trials were submitted to advanced yield trial (AYT). These genotypes were WAB 95 B‐B‐40‐HB (the best performing line among lines received through STRASA and two best lines selected from Upland Regional performance trial (ART3‐11L1P1‐B‐B‐2 and ART8‐L15P14‐1‐2‐1) as well as three genotypes that performed well among 600 new lines developed at Namulonge. The 2011 season II was suitable for selecting high yielding diseases resistant. For instance, WAB95‐B‐B‐40‐HB and WAB788‐16‐3‐2‐1‐HB earlier selected had considerable symptoms of BLS and narrow leaf spot.
Please check the section heading "2.5. Variety and status of deployment" and also its subsequent text is incomplete.
In the year 2013, six best performing rice varieties were presented for release to the Variety Release Committee in Uganda. Among the traits and Characteristics that was provided as evidence of superiority to the existing rice varieties were, higher yield, preferred grain and cooking qualities, maturity, tolerance to stresses especially drought.
A list of only 30 genotypes, including top 20 and bottom 10 least performing genotypes, in terms of filled grains are presented in Table 3. Among the top 20 genotypes, three namely NERICA 7, CO 39, and VANDANA were reference materials for high drought tolerance at reproductive growth stage. There were nine out of the 20 lines from the CT breeding lines and five from the NERICA generations.
No. | Genotypes | Filled grains (%) |
---|---|---|
Top 20 genotypes | ||
112 | WAB 56‐50 | 96.3 |
53 | CT 16333(1)‐CA‐18‐M | 91.1 |
34 | CT 16326‐CA‐3‐M | 89.1 |
101 | NERICA 14 | 88.7 |
108 | WAB 56‐39 | 88.6 |
137 | NERICA 7 | 88.1 |
132 | CO 39 | 87.8 |
142 | VANDANA | 87.6 |
124 | NERICA 6 | 87.4 |
83 | CT 16340‐CA‐9‐M | 86.7 |
190 | NERICA 17 | 86.6 |
45 | CT 16329‐CA‐10‐M | 85.5 |
177 | WBK 35 (F3) | 84.9 |
92 | CT 16315(1)‐CA‐1‐M | 84.7 |
1 | CT 16330(1)‐CA‐2‐M | 84.1 |
165 | IR 64 | 83.8 |
188 | NERICA 15 | 83.5 |
90 | CT 16307‐CA‐5‐M | 83.5 |
10 | CT 16353‐CA‐17‐M | 83.4 |
30 | CT 16324‐CA‐10‐M | 83.1 |
Bottom 10 genotypes | ||
169 | IR 57514‐PMI 5‐B‐1‐2 | 49.4 |
80 | CT 16316‐CA‐2‐M | 49.4 |
106 | IDSA 6 | 49.1 |
104 | ITA 123 (FKR 28) | 47.9 |
175 | RAM 118 | 47.8 |
49 | CT 16346‐CA‐11‐M | 47.8 |
32 | CT 16312(1)‐CA‐1‐M | 47.3 |
166 | IR 77298‐14‐1‐2 | 45.7 |
155 | LAC 23 | 43.8 |
65 | CT 16307(1)‐CA‐2‐M | 27.9 |
Overall | Mean | 67 |
LSD0.05 | 1.88 | |
CV% | 13.2 | |
Range/LSD | 17.6 | |
Variance | 18.6 |
The top 20 and bottom 10 genotypes in terms of percent filled grains.
Generalized linear analysis for different traits pooled across sets and sites are presented in Table 4. Results showed that both GCA and SCA effects within sets for filled grains, grains per panicle, leaf area tiller number, and number of panicles per plant were significant (P = 0.001), while only the GCA effects within sets for tiller number were significant (P = 0.001) but not the SCA effects.
Source of variation | Mean square value | ||||||
---|---|---|---|---|---|---|---|
d.f | Spikelet fertility | Grains per panicle | Leaf area | Plant height | Tiller number | Panicle number | |
Env1 | 1 | 146.5*** | 1441.2*** | 1335.9*** | 109.3*** | 1534.8*** | 1708.8*** |
Set | 1 | 41.5*** | 13.6*** | 35.0*** | 10.1*** | 0.7 | 14.1*** |
Set/GCAf | 4 | 10.1*** | 10.5*** | 17.3*** | 7.3*** | 3.2** | 5.1*** |
Set/GCAm | 4 | 5.8*** | 11.4*** | 12.8*** | 26.5*** | 8.3*** | 3.8** |
Set/SCA | 8 | 3.6*** | 8.5*** | 8.3*** | 26.5*** | 1.2 | 14.6*** |
Env × Set | 1 | 14.9*** | 15.4*** | 35.4*** | 1.9 | 6.3** | 13.2*** |
Env × Set/GCAf | 4 | 14.8*** | 8.4*** | 14.7*** | 1.4 | 4.5*** | 0.3 |
Env × Set/GCAm | 4 | 5.6*** | 7.9*** | 11.9*** | 3.7** | 2.5** | 0.1 |
Env × Set/SCA | 8 | 2.8** | 7.2*** | 8.3*** | 1.8 | 1.6 | 0.5 |
Pooled mean square for filled grains and other secondary traits under drought stress and nondrought stress \x3c!--
* P < 0.05.\x3c!-- Please check the section hierarchy and its numbering and amend if necessary.
** P < 0.01.
*** P < 0.0010.
1 Environment.
The male and female mean squares were all significant (P = 0.05) for the filled grains under drought stress (DS) and nondrought stress (NDS) conditions for the A set population (Table 5). There was significant (P < 0.05) mean square for male × female interaction for the filled grains under NDS for set A. In the case of the B crossing set, the male × female interaction mean squares were significant under DS and NDS conditions. In addition, the mean square of male and female were significant under DS but not under NDS. The male, female, and male × female interaction mean squares were all highly significant (P = 0.001) for the total number of grains per panicle under NDS conditions for the A set and significant (P = 0.05) under DS conditions. In the case of B crossing set, male, female, and the male × female interaction, mean squares were significant under NDS conditions, but not the case under DS conditions. The set A had highly significant (P < 0.001) male, female, and male × female mean squares for leaf area under NDS conditions. Mean squares for male and male × female interactions were significant for leaf area under DS conditions, but not the case of female mean square. The results of the B crossing set revealed that male, female, and the male × female interaction mean squares were all highly significant (P < 0.001) under the NDS and DS conditions.
Mean square values for sets A and B | |||||
---|---|---|---|---|---|
Set A | Set B | ||||
Source | d.f | Drought stress | Nondrought stress | Drought stress | Nondrought stress |
Fertility1 | |||||
Male | 2 | 3.21* | 12.90* | 0.51* | 1.86 |
Female | 2 | 4.07* | 9.01* | 2.58* | 5.40 |
Male × female | 4 | 1.56 | 6.84* | 3.29* | 0.80** |
Total grains per panicle | |||||
Male | 2 | 4.58* | 8.60*** | 2.39 | 12.28*** |
Female | 2 | 4.14* | 9.86*** | 1.38 | 9.51*** |
Male × female | 4 | 2.64* | 9.40*** | 0.62 | 5.83*** |
Leaf area | |||||
Male | 2 | 3.09* | 11.00*** | 6.96*** | 15.51*** |
Female | 2 | 1.40 | 16.21*** | 10.01*** | 16.01*** |
Male × female | 4 | 4.76*** | 9.74** | 17.74*** | 5.34*** |
Plant height | |||||
Male | 2 | 4.46* | 4.45 | 26.70*** | 26.44*** |
Female | 2 | 1.26 | 1.20** | 7.66*** | 7.27*** |
Male × female | 4 | 8.14*** | 8.14*** | 15.70*** | 14.70*** |
Tiller no | |||||
Male | 2 | 23.13* | 42.99 | 26.70*** | 4.61** |
Female | 2 | 3.36 | 49.08 | 7.66*** | 0.75 |
Male × female | 4 | 2.17***[23] | 13.97 | 15.70*** | 1.67 |
Mean squares for filled grains, total grains per panicle, leaf area, plant height, and tiller number under drought and nondrought stress.
* P < 0.1,
** P < 0.05,
*** P < 0.001
1 Filled grains in percentage.
The male × female interaction mean squares were all highly significant (P = 0.001) for the plant height under DS, and NDS conditions for the A and B populations. There was significant mean square for female effects under NDS, but not under DS conditions for the A and B populations. In the case of B crossing set, male, female, and the male × female interaction mean squares were all highly significant under both NDS and DS conditions. Mean squares for male, female, and male × female interaction mean squares were not significant for the tiller number under NDS conditions for the A populations. There was, however, significant mean square for male and the male × female interactions but not female mean squares under DS conditions. On the other hand, the B crossing set had male, female, and the male × female interaction mean squares, all highly significant, under both DS and only the male under NDS conditions.
General combining ability (GCA) for female and male (GCAf and GCAm) in A populations under DS and NDS conditions are presented in Figure 1. The total GCA for both male and female parents (GCAt) under DS were more than 55% for all five traits except leaf area that had 42%. All the SCA values were less than 50%. The SCA effects of tiller number, under DS, was 14% and the male × female interaction was not significant (Table 5). Similarly, the SCA effects for filled grains were not important under DS conditions because there were lack of significance (Table 5). This finding, therefore implies that the additive effects was more important than nonadditive effects for filled grains, total number of grains per panicle, plant height, and tiller number.
Relative (%) contribution of GCA and SCA effects to the cross sum of squares in set A under drought stress and nondrought stress.
Under NDS, however, filled grains, leaf area, and tiller number had GCAt more than 55%. The total number of grains per panicle and the plant height had nearly equal GCAt, when compared with SCA. This finding implies that the additive effects are more important than nonadditive effects for filled grains, leaf area, and tiller number, while additive and nonadditive effects had nearly equal effects for total number of grains per panicle. However, the lack of significance in the male × female interactions for tiller numbers (Table 5) makes the importance of SCA not valid.
\nResults of the GCA effects for female and male (GCAf and GCAm) for B populations, under DS and NDS conditions, are shown in Figure 2. The GCA total (GCAt) under drought was more than 55% for filled grains, total number of grains per panicle, and tiller number under DS. The GCAt and SCA for plant height were nearly equal, while a very high SCA value of 68% was found for leaf area. All the SCA values were less than 50%; moreover, the SCA effects for tiller number were not significant (Table 5) and that of total number of grains per panicle were also not significant (Table 5). This finding implies that the additive effects were more important than nonadditive effects for filled grains, total number of grains per panicle, and tiller number, while additive and nonadditive effects had nearly equal effects for plant height. Under NDS conditions, however, filled grains, total number of grains per panicle, leaf area, and tiller number had GCAt more than 55%. The plant height had nearly equal GCAt when compared with the SCA. This finding implies that the additive effects were more important than nonadditive effects for filled grains, total number of grains per panicle, leaf area and tiller number, while additive and nonadditive effects had nearly equal effects for plant height.
Relative (%) contribution of GCA and SCA effects to the cross sum of squares in set B under drought stress and nondrought stress.
Table 6 showed the GCA effects for filled grains for interspecific and intraspecific rice. The GCA values for filled grains were the only one presented, because other secondary traits had weak correlation with the filled grains, which is a trait associated with drought tolerance. Positive GCA effect is desirable in breeding for improved drought tolerance. Strong negative values of GCA effects of parents show contribution of GCA towards low filled grains, while high positive values show high filled grains. Since both GCA effects and SCA effects were significant for filled grains, the individual values for both GCA and SCA effects are presented (Tables 6 and 7). Parents CT 16350‐ CA‐5‐M, IRAT 325, and NERICA 9 had positive and significant scores of filled grains under NDS conditions. In the DS environment, CK 73 was highly positive and significant at P = 0.001, while CT 16344‐CA‐9‐M, NERICA 9, and CT 16346‐CA‐20‐M had positive and significant filled grain scores at P = 0.01.
Filled grains | ||
---|---|---|
Nondrought stress | Drought stress | |
Female | ||
NERICA 8 | -0.46 | -3.42** |
NERICA 13 | -1.69 | 3.45** |
IRAT 325 | 2.15* | -0.03 |
Bonanca | -2.80** | 1.04 |
WITA 1 | -4.98*** | -6.23*** |
CK 73 | -2.18* | 5.19*** |
Male | ||
CT 16334(2)‐CA‐2‐M | -1.29 | -1.18 |
WAB 365‐B‐1H1‐HB | -0.71 | -2.12* |
NERICA 9 | 2.00* | 3.30** |
CT 16346‐CA‐20‐M | -2.54** | -2.26* |
CT 16350‐ CA‐5‐M | 5.26*** | -1.16 |
CT 16344‐CA‐9‐M | -2.72** | 3.42** |
SE | ±0.83 | ±0.97 |
Estimates of general combining ability (GCA) effects for filled grains under drought and nondrought stress conditions.
* Significant at 0.05 (2.15).
** Significant at 0.01 (2.98).
*** Significant at 0.001 (4.14).
Filled grains | ||
---|---|---|
Nondrought stress | Drought stress | |
NERICA 8 × CT 16334(2)‐CA‐2‐M | -3.46 | -12.86*** |
NERICA 13 × CT 16334(2)‐CA‐2‐M | 1.47 | 4.48* |
IRAT 325 × CT 16334(2)‐CA‐2‐M | 0.73 | 7.21** |
NERICA 8 × WAB 365‐B‐1H1‐HB | -0.59 | 8.53*** |
NERICA 13 × WAB 365‐B‐1H1‐HB | -2.01 | -4.93* |
IRAT 325 × WAB 365‐B‐1H1‐HB | 1.90 | -5.71* |
NERICA 8 × NERICA 9 | 3.60 | 0.92 |
NERICA 13 × NERICA 9 | -1.12 | 3.91 |
IRAT 325 × NERICA 9 | -0.46 | -1.52 |
Bonanca × CT 16346‐CA‐20‐M | 6.15** | -3.19 |
WITA 1 × CT 16346‐CA‐20‐M | -6.39** | -0.88 |
CK 73 × CT 16346‐CA‐20‐M | -2.32 | 1.86 |
Bonanca × CT 16350‐ CA‐5‐M | 1.30 | -1.74 |
WITA 2 × CT 16350‐ CA‐5‐M | 7.12** | -1.63 |
CK 73 × CT 16350‐ CA‐5‐M | -3.17 | 2.26 |
Bonanca × CT 16344‐CA‐9‐M | -10.27*** | 6.02** |
WITA 2 × CT 16344‐CA‐9‐M | 4.24* | -3.67 |
CK 73 × CT 16344‐CA‐9‐M | 3.30*** | 1.12 |
Estimates of specific combining ability (SCA) effects for filled grains under drought and nondrought stress conditions.
* Significant at 0.05.
** Significant at 0.01.
*** Significant at 0.001.
Superior crosses were observed, with positive SCA effects (Table 7). Under nondrought stress conditions, crosses WITA 1 × CT 16350‐ CA‐5‐M, Bonanca × CT 16346‐CA‐20‐M, and CK 73 × CT 16344‐CA‐9‐M had significant filled grain score of 0.01%, 0.01%, and 0.001%, respectively. The cross WITA 1 × CT 16344‐CA‐9‐M had significant filled grain score at 0.05 level of significance. In the drought stress conditions, the cross NERICA 8 × WAB 365‐B‐1H1‐HB were highly significant at P = 0.001, and Bonanca × CT 16344‐CA‐9‐M and IRAT 325 × CT 16334(2)‐CA‐2‐M were positive and significant at 0.01.
The mean, variance, and mean variance of filled grains for P1, P2, F1, F2, and F3 are shown in Table 8. The F2 populations had the highest variance followed by F3 and F1. Scaling tests for dominance × dominance and additive × additive interactions were nonsignificant for both levels. Dominance main effects were not significant, but additive main effects were significant at P = 0.01. When the mean scores were fitted to an additive model, it fitted with r2 = 0.77 (Figure 3). Mean filled grains score was best linear unbiased estimator (BLUE) of the traits
Descriptive summary of generations | ||||
---|---|---|---|---|
Generations | d.f | Mean | Variance | Mean variance |
P1 | 29 | 77.80 | 42.92 | 2.59 |
P2 | 29 | 56.93 | 16.89 | 1.90 |
F1 | 29 | 74.73 | 80.47 | 2.49 |
F2 | 59 | 72.07 | 141.08 | 1.20 |
F3 | 59 | 64.60 | 81.87 | 1.08 |
Scaling test for filled grains | ||||
Interactions | Scale | SE | d.f | t (Scale/SE) |
dominance × dominance | -15 | 3.559 | 146 | -1.184NS |
additive × additive | 3 | 3.361 | 176 | -0.893NS |
Components of means (three parameters) | ||||
Gene effects | Expectation estimates | SE | t = (component/SE) | d.f |
Mean | 57.0 | 1.176 | 48.46** | 59 |
Additive effects | 10.5 | 0.707 | 14.85** | 58 |
Dominance effects | 0.8 | 36.842 | 0.22 | 147 |
Summary of generations in variety 18 × 138 cross, scaling test, and components of means for filled grains score.
Proportions of genes contributing to filled grains score.
The estimate of the number of genes that control filled grains trait based on Castle‐Wrights method was 0.9 ≈ 1 gene. Estimate of the degree of dominance in the F1 and F2 generation based on the [15] method was -3 ≈ 0 and 0.9 ≈ 1 level of dominance, respectively.
\nThe narrow sense heritability in the generations from the cross between CT 16334 (2)‐CA‐2‐M and WAB 450‐1‐BL1‐136‐HB using regression of F1 on mid‐parents and F2 to F1 based on single seed decent are shown in Figures 4 and 5 respectively. In the F1 to midparent regression, heritability of 60% was realized, but when F2 was regressed onto F1 means, the heritability estimate was 74%.
Regression of F1 progenies on midparents for 12 × 138 cross using filled grains.
Regression of F2 progenies on F1 parental means for 12 × 138 cross using filled grains.
Results of evaluation of two sets of new 660 at NaCRRI are presented in this section. The first set grown under optimum moisture throughout the growth period is presented in Table 9. The selection pressure was 11.4% (75 out of 660 rows selected) for rain‐fed lowland conditions and 9.85% (65 out of 660 rows selected) for rain‐fed upland conditions. Candidate varieties CAIAPO/CT 16324‐CA‐9‐M, WAB 450‐1‐BL1‐136‐B/WAB 450‐B‐136‐HB, CT 16317‐CA‐4‐M/WAB 365‐B‐1H1‐HB, IRAT 325/WAB 450‐B‐136‐HB, and CT 16342‐CA‐25‐M/CK 73 are among the lines. Overall, 84 genotypes were selected for further evaluation.
Selection under rain‐fed upland conditions | Selection under rain‐fed lowland conditions | ||||
---|---|---|---|---|---|
No | Groups of crosses | Total | No | Groups of crosses | Total |
One row crosses selected at F4 | One row crosses selected at F4 | ||||
1 | Bonanca × WAB 881‐10‐37‐18‐3‐P1‐HB | 1 | WAB 56‐104 × CT 16324‐CA‐9‐M | ||
2 | IRAT 325 × WAB 450‐B‐136‐HB | 2 | CT 16350‐ CA‐5‐M × WITA 2 | ||
3 | CT 16355‐CA‐15‐M × IRAT 112 | 3 | CT 16355‐CA‐15‐M × IRAT 112 | ||
4 | WAB 365‐B‐1H1‐HB × WAB 450‐1‐BL1‐136‐HB | 4 | CT 16317‐CA‐4‐M × IRAT 104 | ||
5 | WAB 450‐B‐136‐HB × IRAT 325 | 5 | WAB 450‐B‐136‐HB × IRAT 325 | ||
6 | CT 16344‐CA‐9‐M × WITA 2 | 6 | WAB 365‐B‐1H1‐HB × IRAT 325 | ||
7 | WAB 365‐B‐1H1‐HB × IRAT 325 | 7 | CT 16313‐CA‐4‐M × Caiapo | ||
8 | WBK 35 (F3) × WAB 450‐1‐BL1‐136‐HB | 8 | WAB 56‐104 × CT 16313‐CA‐4‐M | ||
9 | Bonanca × CT 16346‐CA‐20‐M | 9 | CK 73 × CT 16346‐CA‐20‐M | 9 | |
10 | CT 16342‐CA‐25‐M × IRAT 257 | Two rows crosses selected at F4 | |||
11 | WAB 450‐B‐136‐HB × WAB 365‐B‐1H1‐HB | 1 | IRAT 13 × CT 16342‐CA‐25‐M | ||
12 | CT 16334(2)‐CA‐2‐M × IRAT 325 | 2 | CT 16346‐CA‐20‐M × Bonanca | ||
13 | CT 16344‐CA‐9‐M × CK 73 | 3 | CT 16342‐CA‐25‐M × CK 73 | ||
14 | CT 16344‐CA‐9‐M × Bonanca | 4 | WAB 365‐B‐1H1‐HB × IRAT 325 | ||
15 | Bonanca × WAB 450‐I‐B‐P‐38‐HB | 15 | 5 | IRAT 112 × WAB 365‐B‐1H1‐HB | 10 |
Two rows crosses selected at F4 | Three rows crosses selected at F4 | ||||
1 | Caiapo × CT 16324‐CA‐9‐M | 1 | Bonanca × WAB 881‐10‐37‐18‐3‐P1‐HB | ||
2 | CT 16313‐CA‐4‐M × WAB 56‐104 | 2 | WAB 365‐B‐1H1‐HB × IRAT 325 | 6 | |
3 | IRAT 325 × WAB 365‐B‐1H1‐HB | Four rows crosses selected at F4 | |||
4 | CT 16324‐CA‐9‐M × WAB 56‐104 | 1 | Caiapo × CT 16324‐CA‐9‐M | ||
5 | WAB 365‐B‐1H1‐HB × IRAT 325 | 2 | WAB 450‐1‐BL1‐136‐HB × WAB 450‐B‐136‐HB | ||
6 | WAB 450‐B‐136‐HB × IRAT 112 | 3 | CT 16324‐CA‐9‐M × WAB 56‐104 | 12 | |
7 | CT 16334(2)‐CA‐2‐M × IRAT 325 | 14 | Five rows crosses selected at F4 | ||
Three rows crosses selected at F4 | 1 | IRAT 325 × WAB 450‐B‐136‐HB | 5 | ||
1 | WAB 450‐1‐BL1‐136‐HB × WAB 450‐B‐136‐HB | Six row crosses selected at F4 | |||
2 | CT 16317‐CA‐4‐M × WAB 365‐B‐1H1‐HB | 1 | WAB 450‐B‐136‐HB × WAB 365‐B‐1H1‐HB | 6 | |
3 | IRAT 112 × WAB 365‐B‐1H1‐HB | Seven rows crosses selected at F4 | |||
4 | CT 16334(2)‐CA‐2‐M × WAB 450‐1‐BL1‐136‐HB | 1 | WAB 365‐B‐1H1‐HB × WAB 450‐1‐BL1‐136‐HB | ||
5 | WAB 56‐104 × CT 16313‐CA‐4‐M | 2 | Bonanca × CT 16346‐CA‐20‐M | 14 | |
6 | CK 73 × CT 16350‐CA‐5‐M | Thirteen rows crosses selected at F4 | |||
7 | IRAT 257 × CT 16355‐CA‐15‐M | 21 | 1 | CT 16317‐CA‐4‐M × WAB 365‐B‐1H1‐HB | 13 |
Five rows crosses selected at F4 | |||||
1 | IRAT 112 × WAB 450‐B‐136‐HB | ||||
2 | IRAT 13 × CT 16342‐CA‐25‐M | ||||
3 | CT 16342‐CA‐25‐M × IRAT 13 | 15 | |||
Rows selected | 65 | 75 | |||
Total Hills planted | 660 | 660 | |||
Selection pressure | 9.85 | 11.4 |
Selection of F4 genotypes from 660 F3 genotypes.
\x3c!-- The paragraph `Results of evaluation of 84...’ is repeated. Similarly, a few sentences are also repeated in the chapter. Please check.
No | Genotype | RYMV | Blast | BLB |
---|---|---|---|---|
1 | P 22 H13 WAB 450‐1‐BL1‐136‐HB × WAB 450‐B‐136‐HB | v | 0 | 0 |
2 | P 36 H1 WAB 365‐B‐1H1‐HB × WAB 450‐1‐BL1‐136‐HB | 0 | 0 | 0 |
3 | 16‐16 CT 16344‐CA‐9‐M × Bonanc | 0 | 0 | 0 |
4 | 13‐13 CT 16344‐CA‐9‐M × CK 73 | 0 | 0 | 0 |
5 | NERICA 4 | 0 | 0 | 0 |
6 | P 25 H1 CT 16346‐CA‐20‐M × Bonanca | 0 | 0 | 0 |
7 | P 8 H2 Caiapo × CT 16324‐CA‐9‐M | 0 | 0 | 0 |
8 | 77 WAB95‐B‐B‐40‐HB | 0 | 0 | 0 |
9 | 96 WAB56‐77 | 0 | 0 | 0 |
10 | 152 AB788‐16‐3‐2‐1‐HB | 0 | 0 | 0 |
11 | P 24 H8 IRAT 13 × CT 16342‐CA‐25‐M | 0 | 0 | 0 |
12 | P 1 H14 Bonanca × WAB 881‐10‐37‐18‐3‐P1‐HB | 0 | 0 | 0 |
13 | P 4 H6 CT 16350‐CA‐5‐M × WITA 2 | 0 | 0 | 0 |
14 | P 29 H1 CT 16342‐CA‐25‐M × CK 73 | 0 | 0 | 0 |
15 | P 23 H1 CT 16346‐CA‐20‐M × WITA 2 | 0 | 0 | 0 |
16 | P 45 H15 WAB 365‐B‐1H1‐HB × IRAT 325 | 0 | 0 | 0 |
15 | P 24 H9 IRAT 13 × CT 16342‐CA‐25‐M | 0 | 0 | 0 |
18 | P 27 H10 CT 16317‐CA‐4‐M × WAB 365‐B‐1H1‐HB | 0 | 0 | 0 |
19 | P 5 H2 IRAT 325 × WAB 450‐B‐136‐HB | 0 | 0 | 0 |
20 | P 29 H4 CT 16342‐CA‐25‐M × CK 73 | 0 | 0 | 0 |
List of 20 varieties that was resistant to RYMV, blast, and BLB in five locations: Namulonge, Kigumba, Kibaale, Lira, and Doho.
Rank | Acc no. | Yield | Rank | Acc no. | Yield | Rank | Acc no. | Yield | Rank | Acc no. | Yield |
---|---|---|---|---|---|---|---|---|---|---|---|
1 | P27‐H14 | 11,950 | 22 | P35‐H5 | 6625 | 43 | P36‐H16 | 6075 | 64 | P22‐H3 | 5156 |
2 | P29‐H4 | 9750 | 23 | P59‐H13 | 6625 | 44 | P5‐H14 | 6025 | 65 | P55‐H9 | 5139 |
3 | P36‐H17 | 9313 | 24 | P59‐H19 | 6625 | 45 | P1‐H14 | 6000 | 66 | P27‐H9 | 5100 |
4 | P5‐H11 | 9111 | 25 | P27‐H15 | 6550 | 46 | P55‐H2 | 5975 | 67 | P27‐H12 | 5071 |
5 | P36‐H9 | 8688 | 26 | P36‐H4 | 6500 | 47 | P31‐H3 | 5950 | 68 | P28‐H3 | 5025 |
6 | P36‐H4 | 8417 | 27 | P55‐H17 | 6500 | 48 | P27‐H10 | 5900 | 69 | P5‐H3 | 4825 |
7 | P51‐H17 | 8400 | 28 | P5‐H2 | 6500 | 49 | P26‐H17 | 5700 | 70 | P8‐H10 | 4594 |
8 | P33‐H3 | 7625 | 29 | P50‐H1 | 6469 | 50 | P59‐H9 | 5700 | 71 | P26‐H13 | 4500 |
9 | P22‐H6 | 7600 | 30 | P1‐H17 | 6400 | 51 | P59‐H8 | 5675 | 72 | P7‐H2 | 4300 |
10 | P25‐H14 | 7600 | 31 | P22‐H13 | 6375 | 52 | P49‐H3 | 5625 | 73 | P45‐H15 | 4275 |
11 | P37‐H13 | 7575 | 32 | P59‐H17 | 6375 | 53 | P33‐H6 | 5583 | 74 | P24‐H9 | 4250 |
12 | P31‐H15 | 7250 | 33 | P26‐H6 | 6350 | 54 | P36‐H1 | 5583 | 75 | P27‐H3 | 4179 |
13 | P34‐H2 | 7188 | 34 | P27‐H11 | 6350 | 55 | P36‐H8 | 5500 | 76 | P8‐H17 | 3700 |
14 | P25‐H1 | 7125 | 35 | P38‐H15 | 6325 | 56 | P27‐H1 | 5464 | 77 | P55‐H19 | 3650 |
15 | P33‐H1 | 7000 | 36 | P7‐H19 | 6325 | 57 | P26‐H18 | 5450 | 78 | P1‐H20 | 3500 |
16 | P26‐H1 | 6889 | 37 | P55‐H5 | 6275 | 58 | P5‐H4 | 5429 | 79 | P4‐H6 | 3400 |
17 | P27‐H18 | 6850 | 38 | P22‐H16 | 6250 | 59 | P7‐H14 | 5357 | 80 | P27‐H17 | 3300 |
18 | P24‐H8 | 6833 | 39 | P59‐H10 | 6214 | 60 | P29‐H1 | 5333 | 81 | P59‐H17 | 3125 |
19 | P55‐H10 | 6775 | 40 | P35‐H12 | 6188 | 61 | P56‐H19 | 5325 | 82 | P27‐H6 | 2800 |
20 | P8‐H2 | 6708 | 41 | P55‐H20 | 6125 | 62 | P23‐H1 | 5300 | 83 | P27‐H2 | 2679 |
21 | P27‐H7 | 6625 | 42 | P58‐H16 | 6083 | 63 | P8‐H15 | 5194 | 84 | P58‐H11 | 2607 |
Yield of 84 breeding lines screened at Lira.
NB: Yield of NERICA 4 was 5600 tons/ha
Results of evaluation of nine selected lines along with two earlier selected lines and a local check is presented in Table 12. Six lines were selected and presented for release to the National Variety Release Committee of Uganda\x3c!-- Please clarify the sentence " Lines[26] selected and presented for release..."
Genotype | Arua | NaCRRI | Masindi | Soroti | Kibaale | Kanungu | Mean yield | Mean rank | Yield under optimal condition |
---|---|---|---|---|---|---|---|---|---|
1 | 2913 | 2653 | 2906 | 3464 | 2619 | 3284 | 2973 | 7.6 | 3500 |
2 | 2747 | 3026 | 3364 | 3703 | 2802 | 3673 | 3219 | 4.2 | 3600 |
3 | 4093 | 2954 | 3069 | 3984 | 3230 | 3561 | 3482 | 4.2 | 4300 |
4 | 2183 | 2538 | 2887 | 3196 | 2287 | 3187 | 2713 | 9.6 | 4500 |
5 | 3785 | 2351 | 2420 | 3454 | 2731 | 2949 | 2948 | 9.4 | 5800 |
6 | 3399 | 2364 | 2496 | 3369 | 2604 | 2974 | 2868 | 9.6 | 3800 |
7 | 4990 | 2214 | 2072 | 3651 | 3068 | 2773 | 3128 | 9.2 | 3750 |
8 | 2660 | 3086 | 3446 | 3726 | 2809 | 3737 | 3244 | 3.2 | 4013 |
9 | 4532 | 3235 | 3326 | 4305 | 3567 | 3837 | 3800 | 1.4 | 4550 |
10 | 4219 | 2937 | 3030 | 4003 | 3264 | 3540 | 3499 | 4.4 | 3650 |
11 | 3656 | 3023 | 3218 | 3928 | 3121 | 3644 | 3432 | 4 | 3600 |
12 | 3928 | 2080 | 2084 | 3286 | 2606 | 2666 | 2775 | 11.2 | 3780 |
Yield of 12 genotypes in six locations in the country and under optimal conditions.
Index[24]:
1. P5H2 (IRAT 325/WAB 450‐B‐136‐HB‐F6).
2. P29H4 (CT 16342‐CA‐25‐M/CK 73‐F6).
3. P8H2 (Caiapo/CT 16324‐CA‐9‐M‐F6).
4. ART3‐11L1P1‐B‐B‐2 ([WAB56‐104/(WAB56‐104/CG14)]/Moroberekan).
5. P27H1 (CT 16317‐CA‐4‐M/WAB 365‐B‐1H1‐HB‐F6).
6. WAB 95 B‐B‐40‐HB (ITA257/(IDSA6/ROK16)).
7. P24H9 (IRAT 13/CT 16342‐CA‐25‐M‐F6).
8. NERICA‐4.
9. P29H1 (CT 16342‐CA‐25‐M × CK 73–F6).
10. P23H1 (CT 16346‐CA‐20‐M/WITA 2‐F6).
11. ART8‐L15P14‐1‐2‐1.
12. P22 H13 (WAB 450‐1‐BL1‐136‐HB/WAB 450‐B‐136‐HB‐F6).
\x3c!-- Please check whether " P24H9 (IRAT 13/CT 16342-CA-25-M-F6)" in the sentence " Seven genotypes namely..." represents the 7th genotype.
Breeding background, characteristics, and selected agronomic information on six varieties were presented to the variety release committee. These varieties were released based on important characteristics detailed in Table 13. The names proposed and accepted by the variety released committee were NamChe‐1. NamChe‐2, NamChe‐3, NamChe‐4, NamChe‐5, and NamChe‐6.
Variety name | NamChe 1 | NamChe 2 | NamChe 3 | NamChe 4 | NamChe 5 | NamChe 6 |
---|---|---|---|---|---|---|
Year of release | 2013 | 2013 | 2013 | 2013 | 2013 | 2013 |
Local name | NamChe 1 | NamChe 2 | NamChe 3 | NamChe 4 | NamChe 5 | NamChe 6 |
Pedigree | WAB95‐B‐B‐40‐HB | NM7‐8‐2‐B‐P‐11‐6 | NM7‐29‐4‐B‐P‐80‐8 | ART3‐11L1P1‐B‐B‐2 | NM7‐27‐1‐B‐P‐77‐6 | NM7‐5‐2‐B‐P‐79‐7 |
Parents | ITA257/(IDSA6/ROK16) | Caiapo/CT 16324-CA-9-M-F6[25] | CT 16342‐CA‐25‐M/CK 73 | [WAB56‐104/(WAB56‐104/CG14)]/Moroberekan | CT 16317‐CA‐4‐M/WAB 365‐B‐1H1‐HB | IRAT 325/WAB \x3c!-- |
Test names | WAB95‐40 | NM7‐1 | NM7‐8 | ART3‐10 | NM7‐6 | NM7‐7 |
Breeding center | AfricaRice, Senegal | NaCRRI, Uganda | NaCRRI, Uganda | AfricaRice, Ibadan | NaCRRI, Uganda | NaCRRI, Uganda |
Characteristics | ||||||
Leaf planotype | Semi‐erect | Semi‐erect | Erect | Erect | Semi‐erect | Erect |
Culm inclination | Semi‐erect | Semi‐erect | Erect | Erect | Erect | Erect |
Culm length (cm) | 64 | 66 | 66 | 65 | 60 | 62 |
Duration from germination to harvest (days) | 110 | 132 | 125 | 120 | 125 | 125 |
Milling percentage | 66.2 | 68.7 | 72.7 | 72.1 | 71.4 | 70.4 |
Volume expansion | 1.7 | 1.6 | 1.6 | 1.9 | 2 | 1.9 |
Yield (kg/ha) | 3800 | 4300 | 4550 | 4500 | 5800 | 5000 |
1000 grain weight (g) | 29 | 28 | 24 | 27 | 26 | 23 |
Grain length dehusked (mm) | 6.3 | 6.7 | 6.3 | 6.5 | 6.4 | 6.4 |
Grain width dehusked (mm) | 2.4 | 2.2 | 2.2 | 2.1 | 2.2 | 2.2 |
Major characteristics of the released varieties.
Results that nine out of 20 best lines were from the CT breeding work imply there was adequate variability in the selected set for selection of suitable genotypes. There were five out of 20 genotypes also indentified as drought tolerant. Also, result that three reference genotypes namely NERICA 7, CO 39, and VANDANA were suitable for identified as drought tolerant in this study implies that the method of screening was acceptable.
The analysis of F2 crosses revealed the various components of gene action controlling various drought tolerance traits in rice. Both male GCA and female GCA effects were significant for filled grains under both DS and NDS, for the A populations and the B populations under DS. The finding that the additive effects were more important than nonadditive effects for total number of grains filled, grains in set A under NDS and B under NDS and DS, implies that additive effects control the traits in different populations and the nonadditive effects varied with populations under study. This result is contrary to the finding by Mohapatra and Mohanty [16] that filled grains was predominantly controlled by nonadditive gene effects under drought stress. However, in the study by Mohapatra and Mohanty [16], the populations were generated by crossing O. sativa with O. sativa. The mechanism of drought tolerance in O. glaberrima, a parent of the interspecific rice used in the current study, was reported to be different from that of O. sativa [17]. This could explain the apparent differences in findings of the current study when compared with that of Ref. [16]. Based on the current study, breeding methods that involve selection in the early generations are recommended. The methods include single seed decent, pedigree selection, and modified bulk methods. Studies using more populations generated from O. sativa and interspecific rice could confirm our finding that the importance of SCA varies with population under study.\x3c!-- `Akbar et al. (1985)’ was cited in the text, but not provided in the reference list. Please check.
Findings of this study that nonadditive effects for total number of grains per panicle was important in both set A and set B under NDS conditions, implies that breeding methods that involve late selection could improve drought tolerance under NDS using number of grains per panicle trait. The use of yield components including grains per panicle has been demonstrated to be effective in improving yield under drought stress by selecting under NDS conditions [18]. The differences between the responses under DS and NDS conditions for total number of grains per panicle could be due to fewer loci within the set B that could segregate for the trait than in A. Set B comprised of lines with more susceptibility to drought stress than those in set A.
\nAdditive effects for number of grains per panicle were important in all the population in set A and set B, under DS and NDS. This implies that breeding methods that involve selection in the early generations especially, single seed decent, pedigree selection, and modified bulk methods could improve drought tolerance through selection of number of grains per panicle. In another study involving O. sativa parents that included susceptible, moderately susceptible, moderately resistant and resistant lines, number of grains per panicle was reported to be controlled by additive effects under NDS conditions [19]. Genes with additive effects were predominant in the inheritance of number of grains per panicle [16]. Both additive and nonadditive effects were nearly equal in populations in set A, under NDS. These set of populations could be used to improve drought stress using methods that involve selection in the early and late generations of the populations. These methods include modified bulk methods and repeated crossing at the segregation stage. Similarly, additive and nonadditive gene effects were significant for number of spikelets per panicle under both normal and saline conditions, and repeated crossing has successfully been used to improve salinity tolerance [20].
\nFindings of this study that nonadditive effects for leaf area were more important than additive effects in both set A and set B under DS conditions, suggests that late selection could improve drought tolerance. In addition, the findings that additive effects were more important than nonadditive effects for the populations in sets A and B under NDS implied that selection methods that involve early selection could be employed under NDS. In the populations in sets A and B, interspecific rice genotypes generated from O. glaberrima crosses were the majority of the parents. O. glaberrima is known to have high vegetative growth as a drought stress adaptation mechanisms [21, 17]. It is likely that these traits were transmitted to the populations under study and it is expressed more under DS than under NDS conditions.
\nResults of this study that additive and nonadditive effects for plant height were nearly equal with contribution for total GCA, varying between 45 and 55% for both set A and set B under DS and NDS conditions, implied that that breeding methods that involve both early and late selection could be employed in the improvement of drought tolerance using this trait. Modified bulk method of selection method could be appropriate. In another study involving O. sativa parents that included susceptible, moderately susceptible, moderately resistant and resistant lines, and plant height was controlled by additive effects under NDS conditions [19]. In the current study, both additive and nonadditive effects were important when the B generations were tested under DS and NDS conditions. Drought traits were controlled quantitatively.
\nThe current study found that additive effects were the more important in the transmission of drought tolerance using tiller number as evidenced by the lack of significance for male × female interaction effects for tiller number. This finding is contrary to the work reported by other scientists that nonadditive effects were more important under drought stress conditions [22, 23]. In another study, however, expression of tiller number, under both NDS and DS situations, was found to involve nonallelic gene interactions [20].
\nOverall, in situations where nonadditive effects are more important, selection should be delayed until later generations. In these types of populations, repeated crossing in the segregating generations may be useful to pool all the desirable genes in one genotypes according to Ref. [24]. The modified bulk method is another useful method of improvement. However, when additive affects are more important, then a modified pedigree method that involves bulking germplasm before evaluation is appropriate. However, when both additive and nonadditive effects are important, two options can be taken depending on the objective of the breeding and the relative importance of the additive or nonadditive effects. In case, if the objective is to develop hybrid rice, as it is planned in Uganda, then pure line selection should be employed. In this approach, additive effects will be extracted because rice is autogamous [25]. In a situation, where both additive and nonadditive gene action are to be exploited, a modified bulk breeding method would hasten the rate of genetic improvement. Similar exploitation of both additive and nonadditive gene action has been conducted in the improvement of cold tolerance [26] and sodicity tolerance in rice [27].
Generally, there was no clear distinction in combining ability between O. sativa and interspecific rice lines under nondrought stress conditions, but the interspecific lines were better combiners under drought stress conditions. Among the O. sativa line, IRAT 325 was a good general combiner, while CT 16350‐CA‐5‐M and WAB 450‐B‐136‐HB (NERICA 9) were good combiners under nondrought stress conditions. In the drought stress condition, however, CK 73, an O. sativa genotype, was the best combiner for improved filled grains. Other parents with lower levels of significance were CT 16344‐CA‐9‐M, WAB 450‐B‐136‐HB (NERICA 9), and CT 16346‐CA‐20‐M.
\nSpecific combining ability analysis revealed that crosses WITA 1 × CT 16350‐CA‐5‐M, Bonanca × CT 16346‐CA‐20‐M, and CK 73 × CT 16344‐CA‐9‐M were best under NDS condition. The cross CK 73 × CT 16344‐CA‐9‐M had both parents as good combiners indicating additive × additive type of gene action. It is expected that these crosses could provide transgressive segregants that could be selected using pedigree methods [28]. The others crosses had mixed combiners, therefore additive and nonadditive gene action could be the major contributors. In such crosses, bulk breeding methods could exploit both gene actions.
There were significant differences among generations for filled grains indicating the presence of sufficient genetic variability. Variability for various traits of rice has been reported [29–32]. The scaling test showed that additive genetic effects but not dominance and epistatic genetic effects were important in the inheritance of filled grains. Fitting means of filled grains on the additive model showed that additive effects accounted for 77% of the genetic variation. In addition, the finding that dominance level was 0 in the F1 population showed that there were no dominance effects.
\nThe generation means analysis confirmed that additive effects were significant in the transmission of filled grains in the populations generated. This study had no inconsistencies in detecting that additive effects were the most important genetic factor in the population under study. In addition, results where narrow sense heritability was high indicated that a high proportion of genetic components of variance can be fixed in segregating generations. Since the selection was conducted under drought stress, it is appropriate that selection for improved drought stress is conducted as early as F2 in the study location. According to Ref. [31], it is appropriate that selection for improved drought stress is conducted using heritability estimates for target traits. There is limited information on the inheritance of filled grains trait under drought stress. However, various reports indicated that additive effects were the main components that controlled the transmission of this trait under high temperature [33, 34]. A single gene pair was estimated to control filled grains under drought stress. A single gene was found to be responsible for the transmission of filled grains under high temperatures [33, 34].
Results of evaluation of two sets of new 660 genotypes showed that CT lines namely CAIAPO, CT 16324-CA-9- CT 16317-CA-4-M and CT 16342-CA-25-M had the highest number of parents that could improve the landraces. These lines were developed for drought tolerance through CIAT Colombia Breeding program.
\nResults of evaluation of 84 rain-fed genotypes that P27-H14 P29-H4 P36-H17, P5-H1 P36-H9 and P36-H4 were the preferred genotypes based on resistance to diseases and yield concurs with other reports (3, 23) that rice varieties with tropical Japonica have higher resistance to RYMV and other diseases.
Results of evaluation of nine selected lines along with two earlier selected lines and a local check is presented in Table 12. Although seven varieties were more had higher yields than NERICA‐4, only six were presented for release when information on milling and cooking qualities were considered. These genotypes were: 1. P5H2 (IRAT 325/WAB 450‐B‐136‐HB‐F6), 2. P29H4 (CT 16342‐CA‐25‐M/CK 73‐F6), 3. P8H2 (Caiapo/CT 16324‐CA‐9‐M‐F6), 4. ART3‐11L1P1‐B‐B‐2, ([WAB56‐104/(WAB56‐104/CG14)]/Moroberekan), 5. P27H1 (CT 16317‐CA‐4‐M/WAB 365‐B‐1H1‐HB‐F6), and 6. WAB 95 B‐B‐40‐HB (ITA257/(IDSA6/ROK16).
Breeding background, characteristics, and selected agronomic information on six varieties were presented to the variety release committee. These varieties were released based on important characteristics summarized in Table 13. Information from genetic studies during F2 generation guided selection of promising lines from F2 through F6. Subsequently, promising varieties were nominated for National Performance Trials and eventually released. Four new varieties were released namely, NM7‐8‐2‐B‐P‐11‐6 generated from CAIAPO/CT 16324‐CA‐9‐M cross, NM7‐29‐4‐B‐P‐80‐8 (CT 16342‐CA‐25‐M/CK 73), NM7‐5‐2‐ B‐P‐79‐7 (IRAT 325/WAB 450‐B‐136‐HB), NM7‐27‐1‐ B‐P‐77‐6 (CT 16317‐CA‐4‐M/WAB 365‐B‐1H1‐HB), and NM7‐5‐2‐ B‐P‐79‐7 (IRAT 325/WAB 450‐B‐136‐HB). These varieties were assigned release names, where WAB95‐B‐B‐40‐HB was named NamChe‐1 at the release in Uganda and ARICA 5 by the AfricaRice Breeding Task Force. ARICA acronym means advanced rice for Africa, implying that the harmonized names are to be used by all parties involved. Another variety bred by AfricaRice is NamChe‐1 (ARICA‐5) with designation ART3‐11L1P1‐B‐B‐2. Of the six varieties released, four were bred from Uganda with support from Alliance for Green Revolution in Africa (AGRA) and the other two were developed by AfricaRice through the AfricaWide Rice Breeding Task Force with support from Stress-tolerant rice for poor farmers in Africa and South Asia. These were NamChe‐2 (NM7‐8‐2‐B‐P‐11‐6), NamChe 3 (NM7‐29‐4‐B‐P‐80‐8), NamChe 5 (NM7‐27‐1‐ B‐P‐77‐6), and NamChe 6 (NM7‐5‐2‐ B‐P‐79‐7). The acronym NamChe means Namulonge Mchere (Mchere means uncooked rice in Kiswhili rice). In 2015, over 20,000 ha was under production based on figures of direct seed sale by different producers.\x3c!-- Please check if the edits made to sentence " Of the six varieties released..." are appropriate. Amend if necessary.
This research found that there was adequate variability in the rice population studied for secondary traits for drought tolerance namely, leaf roll and filled grains. However, the filled grains were found to be more informative and therefore recommended for further studies. Of the three rice groups O. sativa, interspecific lines, and O. glaberrima, there was high similarity between O. sativa and interspecific lines. This similarity could make crossing easy.
\nThe genetic studies for drought provided information on the gene action for drought tolerance at reproductive stage of crosses between interspecific and O. sativa genotypes. Evidence of additive, nonadditive, additive × additive, and dominance effects were found for drought stress at reproductive stage. Additive effects were the most important components that controlled filled grains in most of the populations. This suggests that breeding methods that involve selection in the early generation could therefore be helpful in improving rice for filled grains. These methods include pedigree breeding, pure line selection, mass selection, single seed decent and progeny selection. In a few crosses, however, proportion of filled grains was controlled by nonadditive effects. Methods that involve a delay in selection of genotypes would be appropriate for improving filled grains in these populations. Modified bulk methods of selection are proposed to be employed in this breeding. Tests for magnitude of the gene action for filled grains using additive‐dominance model confirmed that additive gene effects were the most important and additive × additive, as well as, additive × dominance effects were not important. Genotypes O. sativa, namely WITA 1 (O. sativa indica), IRAT 325 (O. sativa japonica), CT 16350‐CA‐5‐M (O. sativa japonica), and WAB 450‐B‐136‐HB (NERICA 9) (interspecific) were good combiners under nondrought stress condition for filled grains. In the drought stress condition, however, CK 73, an O. sativa genotype, was the best combiner for improved filled grains. Specific combining ability analysis revealed that crosses WITA 2 × CT 16350‐ CA‐5‐M, Bonanca × CT 16346‐CA‐20‐M, WITA 2 and CT 16344‐CA‐9‐M were best under NDS condition.
\nFollow up of their performance in countries in the region shows that NamChe‐3 (NM7‐29‐4‐B‐P‐80‐8) and NamChe‐2 (NM7‐8‐2‐B‐P‐11‐6) could be mega variety and a major source of disease resistance. In 2015, over 20,000 ha was under production based on figures of direct seed sale by different producers. This is a success story demonstrating the benefit of collaboration and rigorous breeding in the development of locally adapted rice varieties.
This work was supported by the Government of Uganda, AGRA‐PASS program, and AfricaRice Breeding Task Force program of AfricaRice and University of Kwa‐Zulu Natal, in South Africa.
Please provide publisher location for Ref. [2].
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Please provide volume, page range or doi for Ref.[ 9].
Please provide publisher name and publisher location for Ref. [21].
provide journal title for Ref. [28].
The proprioceptors are sensory receptors that refer to the qualities of movement in the postural dynamics and displacements of the body in space; sense that for a human being is determinant in the perception of himself in a here and now. We call this phenomenon proprioceptive perception, which modulates the emotional states of the individual given the circumstances of the present.
Proprioceptive perception is a complex phenomenon of dynamic character that results from the modulation of phenomena of different orders such as physiological, relational and interpretative. In this chapter we will address the questions of how, in the epigenesis of a human being, proprioceptive perception affects his capacity to become aware of his corporeal existence within his contextual situation with others and with the environment in which they exist, moment by moment; and how the proprioceptive experience modulates -in a present- his emotional states in relation to the immediate interactive context maintaining physiological states of the organism congruent to the present circumstances, conserving in adaptation his life in the space in which it exists.
To answer these questions, we will first situate ourselves from a dynamic systemic perspective that reformulates the concept of effective behavior and cognition that derive from the classical vision of neuroscience and psychology based on the Cartesian reductionist functionalist paradigm. This last one understands cognitive phenomena from a representational perspective, where cognition is conceived as an information processing that results in a faithful representation of an external world that operates independently of the organism that perceives it. This has kept science in search of an understanding of the principles and laws of an objective external world, which explains why in the study of perception, there is a prevalence over the exteroceptive senses of an individual (vision, hearing, touch, smell and taste), ignoring the incidence in the sensory integration of the proprioceptive and interoceptive senses. In second place, we will approach how in the origin of the first living organisms are constituted the generative sensorimotor mechanisms of the movements of the effective behaviors that reveal the knowledge of the living beings, to know that it is source and origin of the way of knowing proper of the Homo sapiens: the reflection. Next, we will explain how the synchronization between attentional reflexive movements and corporal movements gives origin to the proprioceptive perception that makes possible the differentiation of the external and internal space of the individual. And following with the phenomena of the emotions, we will explain the characteristics of the ways of moving of a human being and how the proprioceptive perception influences the modulation of these in relation to the conservation of the well-being of the organism in its structural coupling with the environment. It explains the concept of emotional plasticity and the type of practices that restore it, showing evidence of its effectiveness.
We will begin with the concepts that articulate the reflexive logic of this study that approaches the phenomena of cognition from an evolutionary systemic look that comes from the Theory of Autopoiesis based on the “Theory of the Biology of Knowledge” of Humberto Maturana and Francisco Varela [1], which brings about a radical paradigmatic shift that is produced with the evidence that the internal operation of living beings -in their environment- is of a circular and recursive nature. This implies that the cognitive processes are referred to the changes of the internal states, and not to the changes of the external environment within which it is observed, showing how the world they live in, in a present, is the result of an epigenic process and not of a processing of information captured from the environment. This explains the phylogenetic and ontogenetic origin of the cognitive processes of human beings that - in their relationship with others - give rise to the domain of language, which makes possible their capacity to reflect, and with it, to perceive themselves differentiating proprioceptively from others and from the world of objects that they learn to perceive in their culture.
Both authors define that “knowledge is effective action, that is, operational effectiveness in the domain of existence of the living being” [1], specifying that this domain is constituted, moment by moment, in the physiological operation of the living body in interaction with the environment in which it exists. They show how in organisms with motility this operation of a cognitive nature determines the changes of state of the individual in relation to the conservation of his living identity in a changing environment, and not to changes in the environment, revealing that effective behaviors do not respond to the perception of an image or representation of the state of an external world.
Therefore, we will address the explanation of how the first unicellular organisms of the planet autonomously maintain effective or adequate behaviors for their conservation within the environment from which they arise and with which they maintain a continuous interactional relationship, a fact that reveals their knowledge of how to live in a changing environment, thus showing how the cognitive capacities that, as we will see, result from the physiological operation of the organism in its interaction with its environment, are constituted in the evolution of the species.
This reflection originates with the study of living beings -including human beings- as dynamic autopoietic systems [1]. That is, as living systems that are self-generating, moment by moment, referring to the dynamic organization of molecular relations that constitutes them. This organization remains invariant in a flow of internal structural changes within a changing environment, with which it maintains a continuous interactive relationship of reciprocal nature, preserving the organization that defines its identity as a species.
Thus, in the operation of this organization, a network of interactive relationships between molecular components that produce the components that constitute the - metabolic network - is constituted, moment by moment, maintaining an operation of a circular and recursive nature that generates autonomy, which determines, moment by moment, the appropriate internal states for the conservation of the molecular organization that constitutes them as living beings within the changing environment in which they exist [2]. In this way, the dynamics of this molecular system constitute a physiological operation of cognitive nature, that maintains the orders of the interactional relations between the molecules, making possible the existence of a living unity that differentiates from the environment that it exists in a permanent reciprocal interaction with it.
Therefore, all living bodies, with or without nervous system, are autonomous self-referential beings, with the ability to determine the appropriate behaviors for their living conservation. So, the changes of states of the organism that trigger internal or external disturbances to themselves, are specified by their internal autopoietic operation and not by the changes of the external environment; environment with which it maintains a continuous of recurrent and recursive interactive relations of reciprocal character, that is, bidirectional.
Thus, the reciprocal character of the mutual interactions between the living body/niche generates in the epigenesis of the individuals a congruence or correspondence between the structures of both spaces constituting a structural coupling, in which both spaces -delimited by the edge of the organism- modulate with each other without there being control of one over the other, since each domain specifies the structural changes triggered by the disturbances produced in their mutual interactions, moment by moment.
In this way, in this study, the living body/niche dynamic interactional system is considered as the unit of study of an individual’s behavior; two subsystems that constitute the operational domains of behavior: a) the body domain: constituted in the operation of the physiological dynamics that constitute it, and b) the relational domain: generative of the interactional dynamics that are generated in its operation within the environment in which the individual as a whole exists, an interactional space in which behavior is observed.
“…the phenomena of the structural dynamics of a living system and the phenomena that occur in its interactions in the medium, are phenomena of different kind that occur in phenomenal domains that do not intersect, and cannot be expressed one in terms of the other”. Thus, “[…] behavior of a living system is the interactional and relational dynamics through which a living system realizes its living as a particular kind of organism in its domain of existence […] the structural dynamics of the living systems triggers structural changes in the medium, and at the same time the structural changes that take place in the medium as behavior takes place trigger structural changes in the living system. As living takes place in the continuous conservation of autopoiesis and adaptation by the living system through its behavior, the behavior of the living system operates as the guide in the conservation or loss of the living through the coupling of the structural dynamics of the living system and the medium.” [3].
In this way “what we call behavior when observing changes in the states of the organism in its environment corresponds to the description of the movements of the organism in an environment that we point out.” [1]. This means that this environment does not correspond to the world in which the individual lives.
The implication of evidencing the autonomy of living systems, brings a radical epistemological paradigmatic shift by modifying the conception of living beings, since an autonomous system means that it defines itself through mechanisms of self-organization. Therefore, this characterization of living beings modifies fundamental beliefs of the traditional Cartesian, representational and functionalist paradigm that conceives living systems as heteronomous systems, that is, that they are defined - in their conformation and behaviors - through external mechanisms of control (input–output), therefore their world is treated as if it were independent and represented [4]. In this way, classical science defines behavior as responses to external stimuli, being the environment the one that defines the course of structural changes of the living bodies, thus living bodies have no incidence in their evolutionary transformations, for which they would be heteronomous systems. This is exemplified in the following statement by H. Curtis and N. Barnes:
“The characteristics of the behavior of an organism -its sensitivity to particular stimuli and patterns of response to those stimuli- are the product of natural selection, just as much as the shape of the teeth or the feedback loop that regulates blood pressure. Therefore, natural selection is the force or active agent that determines the course of evolution of the identity of living beings, being these mechanical organisms lacking the autonomy to specify their behavior and structural changes in relation to the conservation of the molecular organization that defines their living identity, and actively specify the niche in which they carry out their living through their behavior, conserving themselves in adaptation in a structural coupling with the environment.” [5].
The reformulation of the generative mechanisms of the effective behaviors leads in turn to a reinterpretation of the concept of cognition, which traditionally has been considered as an information processing in which the sensorial surfaces transduce the stimuli of the environment, sending the information to neuronal structures that process it, generating a representation of the state of the world in which the individual is, from which the system selects the effector motor patterns for the appropriate behaviors to the individual’s situations, which result from phylogenetic and ontogenetic learning of an adaptive nature.
The autopoietic theory originates a reformulation of the generative mechanisms of effective or adequate behavior of living beings, as well as the evolutionary processes that give rise to the diversity of anatomo-physiological structures that define the identity of living species. The autonomy of living bodies makes them an active agent in the transformation of themselves, as well as of the environment in which they are found; a phenomenon that occurs in the epigenesis of living bodies. The evolution of the history of structural links between organisms and their environment generates reciprocal transformations that lead to the maintenance of a congruence in the structural changes of both, a fact that makes possible the conservation in adaptation of the organisms. Traditionally, this congruence has been interpreted as the result of effective behaviors in their adaptation to the environment, which result from responses to the stimuli of an external world that operates independently of the operation of the organism, considering them instructive and unidirectional interactions; thus, in the evolution of the organism, the adaptive behaviors would be determining the structure and identity of the species.
The sequences of movements of an organism in coupling with the environment that are observed in its effective behaviors result from “a very specific correlation coordination between sensory surfaces and motor surfaces, …sensory-motor correlations that originate from the first living beings through metabolic transformations proper to the cellular unit” [1], which in the recurrence and recursiveness of reciprocal interactions with the environment are constituted in sensory-motor learnings that specify ways of interacting in the regularities of changes in the environment, keeping invariant the molecular relations that define the molecular organization that preserves the living unity in adaptation.
By way of example, the feeding behavior of an amoeba about to ingest a protozoon is described by means of the extension of a pseudopod. Pseudopods are expansions or digitations of protoplasm associated to structural changes in the local physicochemical constitution of the cellular membrane. How does the global and unified movement of the animal occur in its structural coupling with an environment in which it is also structurally coupled to it? “The presence of the protozoon generates a concentration of substance in the environment that is capable of interacting with the amoeba’s membrane, triggering changes in protoplasmic consistencies, resulting in the formation of a pseudopod. The pseudopod in turn produces changes in the position of the moving animal, thus modifying the number of molecules in the medium that interact with its membrane. This cycle is repeated, and the sequence of displacements of the amoeba, therefore, is produced through the maintenance of an internal correlation between the degree of modification of its membrane and those protoplasmic modifications that we see as pseudopods, a recurrent and invariant correlation is established between a disturbed or sensory area of the organism and an area capable of producing motor displacements, which maintains invariant a set of internal relations in the amoeba.” [1].
We can see, on the one hand, how the continuous structural coupling of the organism with the environment generates the congruence between the structural changes of both, and on the other hand, how the movements of the organism generate correlations of specific structural changes between sensory and motor surfaces that establish interactive relations of reciprocity. These relationships are not instructive, they generate a continuous structural change in which the change of one is in relation to the change of the other, moment by moment. Thus, the changes in the motor surfaces generate, in turn, changes in the sensory surfaces, sensorimotor dynamics generating permanent movements that are observed in the proper behavior of an organism in its environment, a process that, as we see, does not consist of a process of capturing and processing information from an external world that operates independently of the organism’s operation.
In this way, in the physiological operation of the organism, the modulation of processes of sensorimotor activity is generated, constituting a synchronic coordination of structural changes between local zones of the organism that modulate with each other, resulting in a distributed modulation mechanism from which a state of global activity emerges -of a temporary nature. This global state specifies a coherent and unified cognitive state that determines the behavior of the individual in his/her relationship with the environment; a mechanism in which local changes modulate the state of global activity and, vice versa global states modulate the activity of local areas, without the existence of an external or internal agent or force that controls them. Such mechanisms of sensorimotor coordination are constituted in the operation of every living being with or without nervous system -what varies among species are the types of sensorial and effector structures- of a centralized control or product of an external or internal agent that specifies the states of activity of sensorimotor patterns of the organism, as well as dispenses with the idea of a representation of an external world.
Returning to the behavior of the amoeba in its coupling with the environment in which the protozoon is found, they establish a structural coupling, which from an observer’s perspective, the protozoon constitutes the prey as a result of a feeding behavior. This fact that a western human being perceives, observes and interprets from the distinctions of the world of his culture, has no relation with the intentional behavior of the amoeba, since by the way, the amoeba does not distinguish the protozoon nor has perspective of the changing environment, therefore it does not intentionally go towards it with the purpose of swallowing it and thus feeding itself. This anthropomorphic interpretation hides what occurs inside the animal in its structural coupling with the environment, that is, it hides the physiological operation of a cognitive nature of the organism that specifies its changes of states triggered by the disturbances of the environment, in relation to its previous states, a fact alluded to when characterizing said operation as circular and recursive.
In summary, from this systemic perspective, the sensation-movement state of a living body in its environment, in a present, results from the dynamic activity of the sensory-motor operations that gives rise to a global cognitive state - of a temporary character - that specifies the coordinated and synchronic dynamic movement sequences that constitute the coherence and uniqueness of an effective or adequate behavior for the conservation of the organism in adaptation in the structural coupling with the environment. This was illustrated with the case of the amoeba’s behavior, showing the origins of the generative mechanisms of the effective behavior of living beings with motility, as well as the cognitive processes that result from their co-evolution with the environment in which they exist.
From what has been said before, we can distinguish that the environment in which an observer distinguishes the amoeba, does not correspond to the world that it lives from its sensorimotor dynamics that are referred to its internal operation, which on the one hand means that the environment that it knows is its interiority and on the other hand that these dynamics that specify the movements in their structural coupling with the environment are generative of the lived world, a world without perspective of its changing environment.
We could say that the living of a body arises spontaneously in a generative movement of its knowing, which Maturana expresses strictly by saying: “to live is to know and to know is to live” [1], which from our perspective alludes to a fundamental fact that reveals the mode of existence of every living being, namely, both life and knowledge arise in the same act.
Therefore, the autonomous movement of animals with motility is a key to the understanding of cognition and the phenomenon of perception in human beings, as we will see in the following section the knowledge of the body in its environment is the source and origin of the way of knowing of human beings: reflection. In this regard the biological origin of human knowledge is evidenced by unifying its nature: corporeal-spiritual constitutive of a living unit.
From the world of the biology of knowledge we reach the world of philosophy. The phenomenologist Maurice Merleau-Ponty from his exhaustive studies in the human experience, describes the phenomenology of perception, a study that begins with the conviction that “phenomenology is also a philosophy that re-situates the essences within existence and does not believe that man and the world can be understood only from its factuality” [6]. From his studies of the human perception and behavior, he establishes co-relations between the psychism and physiology that lead him to a reformulation of the classic vision of the body-object, saying: “The union of the soul and the body is not sealed by an arbitrary decree between two external terms; one, the object, the other, the subject. This union is consummated every moment in the movement of existence. It is existence that we find in the body when we approach it through a first way of access, that of philosophy.” [6].
Bearing in mind that the world that a living organism feels within its coupling with the environment, it constitutes a continuum of sensation-movement resulting from the cognitive states that emerge from the dynamics of activity of patterns of sensorimotor correlations. These patterns, which are constitutive of the learning process, result from recurrent and recursive movements that are constitutive of the individual’s behavior, and determine his anatomo-physiological structure, which specifies his species and his way of knowing and living in his structural coupling with the environment.
In the -recurrent and recursive- structural couplings between living beings that co-exist in the same environment, it is constituted temporary reciprocal interactions between them, generating mutual learning that modify in congruence the anatomo-physiological structures of them. For this reason, in each temporary encounter between them the autonomous operating of the corporal structure of each one determines the specific movements of their behavior, recreating the structural couplings that occur in these temporary encounters. For example, this phenomenon occurs with symbiosis relations between species. This is the case with the structural correspondence between pollinating insects and the flowers of the plants they pollinate. The plant species Drakeae glyptodon, an orchid species whose structure takes a similar form to the female Thynnid wasp, and in its operation produces pheromones that attract the male wasp which is the only insect vector of its pollination. Thus, in the co-evolution of both species, they constitute a history of structural coupling that constitutes the structural changes that are conserved in their progeny.
This epigenic phenomenon when it occurs between individuals of the same species gives rise to the constitution of social systems. In the recurrent encounters of two or more organisms, specific action dynamics are synchronically triggered, generating a coordination of action between them resulting in a communication that specifies a particular way of interacting and relating, which defines a domain of possible actions between them. Thus, said systems are constituted in dynamics of networks of coordination of action between individuals, that give emergency to a collective of autonomous beings self-organized, which moves like a totality in congruence to the changes of the environment, inside which the individuals generate behaviors that of isolated form they could not acquire. This is the case of the flocks of Franklin gulls that migrate from Cape Horn to Canada, a flight in which individuals increase their speed of flight by 72%, compared to the speed of flight of an isolated individual, and no further in the case of human beings who, in their social way of life, learn in doing with another to incorporate the mastery of language that makes their capacity for reflection possible.
Thus, in social systems, the learnings that are generated in individuals in the coordination of action among them, constitute the sensorimotor patterns that are the ways of moving and relating that constitute the way of life of the collective, which is transgenerational preserved by maintaining a living knowledge that makes its existence possible within the environment.
This co-evolutionary phenomenon constitutes a communicative process that is not related to an exchange of information, but rather to interactions of a reciprocal nature that generate specific and recurrent structural changes that occur in their encounters; encounters in which the structural changes of the organisms in their reciprocal interactions are modulated - at each instant - generating a coordination of movements that configure a choreography that is repeated in their recurrent interaction within the social system in the environment in which they are found. “We will understand communication as the mutual triggering of coordinated behaviors among the members of a social system” [1].
Following this second order cybernetic perspective [7], which recognizes that the architecture of the neural networks constituting the nervous system that is embedded in the body of the organism, maintains a circular operation that is to say with operational closure, therefore this operation is referred to the states of activity of the network, and not to an external world [8]. This network is self-organized by distributed mechanisms, in which the global states of activity of the network modulates the activity of local zones, and vice versa the activity of local zones modulates the global states of the network. In this way, there is no internal or external agent that controls its operation; on the other hand, this system modulates and is modulated by the physiological operation of the organism. Therefore, the condition of operational closure of the neuronal network would explain that the world perceived by the organism, including the human being, is a world that emerges from the internal operation of the organism in its structural coupling with others and the environment.
Considering this, we will explain the perception of the world lived by a human being and the learnings that originate the proprioceptive perception, from recognizing the type of structural links that occur between the hominid ancestors of Homo sapiens, generative of phylogenetic learnings that make possible the emergence of language and its capacity to reflect. These facts give origin to the particular way of life of a social system constituted by networks of coordinated action coordinations from the operational distinctions of the language domain, that is to say, generative action coordinations of the networks of consensual conversations. We are going to see how the first condition for the constitution of the phenomenon of perception in human beings is the origin of the observer. “The observer and the observed, then, emerge in the flow of structural changes that occur in the members of a community of observers when they coordinate their consensual actions through their recurrent structural interactions in the domain of operational coherences in which they carry out their connected practices of living.” [9]. Thus, if language is constituted in the domain of the coordinations of the dynamics of action coordinations that occur in the joint action of individuals within the social system in which they carry out their living in coexistence.
The mathematician H. Von Foerster, in his original presentation of the notion of second-order cybernetics, who is a precursor of the same, starts by pointing out what he calls a theorem, alluding to the statement made by Maturana after explaining the origin and mode of operation of language, which he states in this way:
“I. - All that is said is said by an observer.
A theorem to which Foerster adds a corollary that affirms:
II.- All that is said is said to an observer”.
Concluding that I and II connect two observers through language, with this connection, in turn, a new concept is established, namely that of society, the two observers constitute the fundamental nucleus of a society, thus the three concepts are connected in a triadic way, each one with the other. “In this interaction we cannot say who was first and who was last […] a closed triad is formed” [7].
Thus, in this circularity, the operation of language within a social system constitutes observers, and observers in turn constitute the language in its operation in the domain of the coordination of the coordination of action among observers, a reflexive dynamic that is recurrent and recursive and generative in turn, of the domain in which it is constituted. This is a dynamic in which perceptual objects that constitute the world of culture in a society of human beings are configured. Thus, the operation of language domain constitutes an operational system constituted by the distinctions of perceptive objects that are associated among them, generating new distinctions that constitute the association network that generates them, a circular and recursive operation.
In the operation of language, the observer and the perceptual objects that constitute him/her are constituted, a circular dynamic that generates the world perceived by individuals, which does not correspond to an objective reality. Therefore, the description that an observer makes of the world of objects or phenomena that he/she perceives is the result of the flow of the experience of his/her consensual behavioral coordination with others. Therefore, these descriptions are not absolute truths, but descriptions agreed upon with others in the coexistence by such “everything said is said by an observer and for an observer” with whom they maintain a generative structural congruence of their coordination of actions in doing together in the coexistence.
“And since perceptual objects arise as behavioral configurations, the world of shared perceptual objects belongs to the sphere of operational concordances between organisms, which constitute them in the course of their coexistence as configurations of their behavioral concordances. In other words, if the perceptual objects remain configured by the behaviors of the organism, the world of perceptual objects that occurs in the coexistence of organisms, including the observer, can only arise from the coexistence as long as the organisms operate generating and conserving their mutual structural correspondence. That this is so is also apparent in everyday life, in which we know that the common world only arises in the community of living” [10].
How does the observer in language generate perceptual objects that are configured in the behavior of the individual? Language occurs in the flow of consensual coordinations of actions of organisms whose actions are coordinated because they have congruent dynamic structures that have emerged or are emerging through their recurrent interactions in a co-ontogenetic structural drift. Because of this, interactions in language are structural interactions that trigger in the organisms interacting contingent structural changes with the course of the coordinations of consensual actions in which they arise. As a result, even though the domain of language is not intercepted with the structural domain of the body of the interacting organisms, the structural changes of the interacting organisms in language are a function of what occurs in their language and vice versa [9].
In this way, the origin of language generates a new operational domain in the behavior of human beings, which generates reflexive operations. Thus, this domain, which is not intercepted with the corporeal domain (constitutive of the physiological operation), nor with the relational domain (constitutive of the reciprocal interactions that the organism maintains with others and its environment), is constituted as a domain that in its operation generates perturbations both in the state of the organism and in the interactive contexts of the organism. Therefore, it corresponds to a third operational domain, which participates in the modulation of human behavior and experience.
The operation of language generates associations, descriptions and interpretations that originate the beliefs of the world of culture that give meaning to the way of doing and relating to individuals, making them learn to incorporate the recurrent and recursive coordination of action. This operational domain, brings the intentional movements from the reflection, which entails the learning of the reflexive movement of orienting the focuses of attention towards the perceptive objects constitutive of the world that the individuals learn to see in the doing with others, within the culture in which they grow. This reflexive attentional movement brings the possibility of the human being to become aware of himself, of the others and of the environment in which they are, by differentiating himself proprioceptively from the objects that he perceives, which occurs by the ways of relating and interacting that are constituted in the way of life of the hominids.
We will now see what happens in the bodily domain of behavior with the structural couplings in the hominid lineages that give rise to the phylogenetic learnings that make the origin of Homo sapiens possible. In the lineages of hominids that give rise to the human being, their way of life was generated learning gave rise to the architecture a nervous system that is characterized by a significant increase in brain mass, which means an increase in interneurons that expands the possibilities of structural plasticity of individuals, and thus the ability to learn, which means greater behavioral plasticity. Today we know that the genomes between Homo sapiens and anthropoids are almost identical, and from neuroscience it is observed that the regions of the brain have equivalents in the brains of apes. An interesting difference is the development of the generative auditory capacity of phoneme learning that is related to the origin of language [11]. Therefore, the advances of science support with their data the assumption that the origin of the reflection capacity of human beings is related to their way of life.
Such learning, which modifies the anatomophysiological structure, occurs in a way of life in which the game generates continuous and recurrent coordinations of action, and in this way increases the capacity to manipulate and differentiate objects with which they interact with others. At the same time, they had daily physical encounters in which they groomed themselves, and caressed in sensual and prolonged interlacing of their bodies and continuous sexual games with prolonged physical contact. In this way these dynamics of action generate structural changes in proprioceptive sensorial surfaces that correlate with modifications in motor surfaces, constituting recurrent changes in the sensoriality of the qualities of the movements of dynamics of postural sequences and positions of the different parts of the body, which together with maintaining a frontal vision with the other in the coordination of movement that are accompanied by guttural sounds, establishing in dynamics of action in which they generate movements of joint generative attentions of a perspective of the movement of itself and the other, which entails the reflective learning to sustain a division of centers of attention in movements of visualization of the movement of the other and proprioceptive sensation of the movements that it coordinates with him. Thus, in these dynamics of structural links of character -recurrent and recursive- learning is produced, which are constitutive of sensorimotor patterns that result from the coordination of visual, proprioceptive, tactile and auditory sensations, which constitute the sensation of movement and space of oneself, which occurs simultaneously with the differentiation of others in space.
This pre-reflective process that is observed in the families of hominids is generating the learning that makes possible in the individuals the reflective movement of coordination of the attentional movements and the corporal movements in relation to one another, generative of a perspective that arises from an internal space delimited proprioceptively towards an external space when dividing its attention. “…in effect, their spatiality is not, like that of external objects or like that of spatial sensations, a spatiality of position, but a spatiality of situation. […] The word here, as it applies to my body, does not designate a certain position with respect to another position or with respect to an external coordinate, but rather the installation of the first coordinates…” [6]. In this way, his corporeality is a spatial reference of his situation in his perspective of the world, moment by moment, which arises in interactions with others and the environment, being the place perceived proprioceptively in which he is and exists in a present.
This reflexive movement constitutive of an observer’s perspective, which arises from learning to divide their focus of attention into movements of joint attention with another observer in the manipulation of their bodies and objects, is configuring perceptual objects that constitute the observer that emerges with them. In this way the reflexive movement that arises from motor couplings between individuals, is constitutive of the operational domain of language and of the structural congruence between the objects of perception and the living body in its structural coupling with the environment: “The observer’s operation in language consists of a way of living in the recursion of behavioral coordination that arise in the community of living and that configure a world of perceptual objects. […] The language and the operation of the observer, therefore, do not require or give rise to references to an external reality. The world of the observer’s descriptions and explanations is a world of modes of coexistence that generate perceptual objects, in which the observer emerges as one of them when language emerges. Hence the generative and transforming power of the world that language and the explanations given in it have.” [12].
This composite concept leads us first to consider how perception is understood from this systemic perspective that includes living beings as structurally determined systems. This means that everything that happens to the organism in the interaction with its environment is determined by its structure. Therefore, the interactions of the organism with its environment are not instructive [1, 12]. From this perspective, perception consists of “the configuration that the observer makes of perceptual objects by distinguishing operational cleavages in the behavior of the organism, by describing the interactions of the organism in the flow of its structural correspondence in the environment” [12].
For there to be a perceptive experience, it is the observer who emerges from language as a perceptual object, the one who configures a world of perceptual objects in the recursion of behavioral coordination that arise in the community of living [13].
On the other hand, we will define proprioception as one more sense like vision, smell, taste, hearing; it is the sense of the qualities of the body’s movement and its situational disposition in space (that the same movements generate) (see Table 1). Proprioception is not in itself a form of perception that gives us the “perception of the body”, it is not the image, nor the representation, nor the consciousness of the body as an object [14]. The proprioceptive sensation is produced, moment by moment, by the changes in the activity of the proprioceptors that generate the dynamics of the postural sequences of the movement of the individual in structural coupling with the environment. This dynamic phenomenon in which the relations of reciprocity between the changes of the sensorial surfaces and the effector surfaces of the movement, generate that the sensation modulates the movement and the movement in turn, modulates the sensation, a continuous flow of sensation-movement. This flow of sensation-movement is constituted in the operation of sensorimotor patterns that specify qualities of behavioral movement. In the reciprocal interactions of the individual in structural coupling with the environment, the cognitive states specified and that in turn specify, the changes in the dynamics of activity selectivity of the sensorimotor pattern networks that give rise to sensory integration (proprioceptive, visual and vestibular) [15] that define the dynamic body scheme, in a present.
Location | Proprioceptor | Quality of sensation |
---|---|---|
Muscle | Spindle afferents Ia & II | Length, speed, acceleration and deceleration Minimal over-contraction force. |
Golgi tendon organ | Dynamic changes of the contraction force | |
Group III y IV | Chemosensitives. Information on metabolic changes and muscle damage/inflammation | |
Joint | Group I & II | Range, speed and position of the joint. Group I (dynamic and static, low threshold, slow adapting), Group II (dynamic, fast adapting) |
Group IV | Feedback on excessive stress on the joint. Sensitive to joint inflammation | |
Skin mechanoreceptors | 5 types of receptors in the skin: two fast adapting and three slow adapting | Contact and texture of objects. The tension of the skin contributes to the sense of movement of the joint. More sensitive to dynamic than static stimulation |
Proprioceptors and quality of sensation.
The body schema is defined as an integrated set of sensorimotor processes that organize perception and action in a non-conscious and sub personal way [16]. The body schema is not phenomenologically available to the observer: “the body schema is not the perception of my body, it is not the image, the representation or even the consciousness of the body. Rather, it is precisely the style that organizes the functioning of the body in communion with its environment [17]. On the other hand, body image includes the immediate conscious perception of the body, including the conceptual construction about the body and the emotional attitude and feelings about the body, “being a complex phenomenon that contemplates at least three aspects: perceptual, cognitive and emotional” [17]. However, other definitions have been proposed for this construct: “cognitive representation of the body based on stored knowledge and sensory experience that underlies perceptual judgments” [18], “a representation of the body’s shape” [19], “perception of the body’s spatial dimension, its size, shape and relative configuration of its parts” [20].
What are we talking about when we talk about proprioceptive perception? Proprioceptive perception differs from the concepts of body schema and body image, since it is a reflexive phenomenon that constitutes an attentional movement of the observer towards the corporeal dimension of his behavior, in a here and now. Thus, proprioceptive perception makes present as object of perception the proprioceptive qualities resulting from the dynamics of postural movement and displacements of the individual in his structural coupling with the environment. These qualities configure the perception of the dynamic corporeal space that is defined in a flow of synchronic coordination of movements of the different parts of the body that configure the coherent and unified global movement constitutive of the proprioceptive qualities that result from the sensorimotor operation of the individual in his structural coupling with the environment.
Thus, proprioceptive perception is the perceptual object of the observer configured with the qualities that make up the internal space that appears sensorially delimited from an external space within which it is situated, generating a perspective of the world of objects from which it differs proprioceptively, perceiving the place in which it exists, in zero time; that is, the living body that constitutes moment to moment, its existence as a living being in a structural coupling with the environment with the capacity to reflect and observe the world that it constitutes in doing with others within its culture.
“If corporeal space and outer space form a practical system, the latter being the background against which it can stand out, or the void before which the object can appear as an objective of our action, it is evidently in the action that the spatiality of the body is carried out, and the analysis of one’s movement has to allow us to understand it better. We understand better, as soon as we consider the body in movement, how it inhabits space (and time, for that matter), because movement is not satisfied with passively supporting space and time, it actively assumes them, it takes them back in their original meaning that is erased in the banality of acquired situations.” [6].
In this way, proprioceptive perception cannot be understood outside of perception-movement. Proprioceptive perception constitutes the reflexive and corporal movements of two dimensions of human behavior constitutive of disjointed operational domains: language and its corporeality. “Reciprocally, every perceptive habit is still a motor habit and here also the capture of a meaning is made by the body.” [6].
So proprioception does not have a dual nature, as proposed by Gallagher [15], since its nature is biological and responds to physicochemical properties. Proprioception corresponds to the body domain; whose operations are the networks of physiological dynamics that constitute the mechanisms of the correlations of the sensory and motor surfaces. While reflection and movements of the focus correspond to the domain of language, whose operations are the networks of semantic distinctions with operational closure. Therefore, when proprioception is a perceptual object of the observer, both the body and language domains are operating simultaneously on proprioceptive perception. In addition, these disjointed domains modulate each other [21], and reflection and attentional movements can trigger changes in proprioception and in turn proprioception generates changes in the language domain, as we will see later.
Consequently, we say that the phenomenon of proprioception is different from the qualities of the perceptual object that the observer configures from his corporeal experience, which results from the modulation of the three operational domains that configure the coherence and uniqueness of his behavior: corporeal, relational and language, moment by moment. Thus, both the proprioception and the proprioceptive perception of the individual in their interactive contexts maintain a structural congruence between both phenomena in their continuous structural changes within their circumstances, thus constituting the effectiveness of their behaviors in relation to both their purposes and the conservation of their well-being.
For this we will first address how muscle physiology is involved in the modulation of body perception. The situational disposition of the individual (his posture and movements, in a present) correlates with a configuration of the afferent activity of the proprioceptors coming from the skin, the joints and the muscles that are projected towards the primary somatosensory cortex and the primary motor cortex, to then converge in higher order somatosensory regions [20]. The integration and comparison of proprioceptive activity with the activity of other sensory modalities (and the reflective capacity of the human being) triggers the perception of the size of the body parts, which is relative to the perception of other body parts, as well as to the environment in which the individual is coupled in a present. Thus, in situations where the activity of the nervous system presents a change in the relationships that are generative of its structure, as is the case of a vascular accident, epilepsy, anesthesia or migraine, the perception of size and shape of body parts will be modulated by this configuration, which is commonly understood as a perceptual “illusion” of the body. This phenomenon has also been observed by applying an external vibration in specific muscle regions [22]. Since the afferent activity of the muscle is modulating the sensation of the position of the limb, when performing such stimulation, it is possible to generate the “illusion” of the perception of the movement of the limb or the whole body in a desired virtual direction.
In these cases, the perception of the body is modified by unintentional factors on the part of the individual. However, the human being, through his reflective capacity, has the ability to direct his attention to the perception of his body and with it modulate the perception of the relative size and shape of his body parts. The evidence shows how paying attention to proprioceptive sensations (directing attention to movement during the execution of a task) generates a change in the sensitivity of the muscle spindle [23, 24], which would be modulating the perception of movement of the individual in its structural coupling with the environment. In this sense, training the proprioceptive perception we can modulate the muscular physiological activity, which as we will see, modulates in turn the sensorimotor correlations of the basic emotions.
In research on emotions, we find a diversity of explanations that involve descriptions of different mechanisms that affect the emotional states of a human being, which respond to different dimensions of the phenomenon: physiological, psychological, relational, behavioral, as well as cultural. Thus, in 1991, Plutchik in his book Emotions [25] indicates more than 57 definitions that arise from various authors in the field of physiology and psychology, such as W. James, S. Freud and B. Skinner, to mention a few. This fact shows the multiplicity of non-linear variables that characterize an emotion, so we can conclude that it is a complex phenomenon, which is naturally observed in the behavior of an individual, and that each person perceives in his experience.
Given this last point, we will understand “emotions as specific sequences of movement of an organism in structural coupling with the environment that an observer distinguishes”. We approach emotional phenomena as the distinction of a specific configuration of a coherence in behavior. In this way we distinguish the phenomena that occur in the different operational domains of behavior: body, relation and language, and correspondingly we observe the correlations of the modes of movement, relationship and interpretation of an individual’s experience.
These specific sequences of movement that constitute modes of movement define possible dynamics of action of the individual in his or her present, and with this the type of interactions that are generated in his or her relational contexts, as well as the distinctions of perceptual objects that originate his or her attentional movements in language, generating his or her interpretations.
In the human being two orders of emotional phenomena are observed that respond to the origin of sensorimotor learning, we find the basic emotions of phylogenetic origin, those -fear, rage, joy and sadness, on which the ontogenetic learning constitutive of the secondary or social emotions are interwoven [26], in the present study only the first ones are approached.
In the basic emotions, patterns of movements generate the activation of specific muscular synergies that are triggered from the autonomic nervous system, and therefore correspond to physiological and cognitive states of the organism. Damasio et al. [27], studied the activity of the central nervous system during the evocation of memories of the 4 basic emotions. In this they observed a specific activation pattern at cortical and subcortical level for each one of the emotions. Furthermore, they observed that the emotional states evoked activate the anterior pontine nucleus, which sends projections to the cerebellum and therefore, would possibly be involved in the activation of specific sensorimotor patterns and the quality of movement of each basic emotion. These findings show that each emotion has a physiological configuration of the nervous system and the motor system that is unique to each state, which correlates with a global cognitive operation that gives rise to the “knowing” of the organism in relation to its environment.
The specific movement sequence patterns we are talking about, correlate with specific sensorimotor patterns that come from a phylogenetic learning, that is, they are sensation-movement patterns that we can identify even in primitive unicellular organisms. Thus, the simple expansion and contraction movements of living bodies are indicative of the approach-avoidance behavioral pattern observable from a cell to the human being [28]. Therefore, from the sensory-motor operation of the organism in its structural coupling with the environment, emerges “the knowing” that is evidenced by the autonomy of the body to determine its effective or adequate behaviors to the maintenance of its living and social identity. That is to say, “knowing” emerges with the minimum living unit that moves and feels, feels and moves constituting the basic emotional movements that preserve the way of being of a living being within an environment that it does not know.
Therefore, these emotions that underlie every secondary emotion are related to the conservation of the individual’s living identity, so that in continuous flow of the changes of state of the organism in its structural coupling with the environment, an emotional state of a cognitive nature can be identified, through the identification of the movements that generate the muscular synergies that are activated autonomously by the physiological operation of the organism. From here we speak of these emotions as a living knowledge that guides our actions in relation to preserving the essential, life.
These basic emotional movements correspond to fear, anger, joy and sadness, which are differentiated by a set of qualities of the sequence of their movements and the activation of muscular synergies [29]. A recent study by Shafir et al. [30], from the analysis of the movement of each one of these emotional states, identified those crucial motor elements that distinguish each emotion and that in turn, in their repetition are capable of evoking an emotional sensation. The results showed that each emotion is predicted by a single set of motor elements and that each motor element is a predictor of a single emotion, suggesting that the 4 emotions under study are discrete and have a biological substrate (see Table 2).
Emotion | Quality of emotional movement |
---|---|
Rage / Anger | Advance with sudden, direct effort. Punching movements and leaning forward. |
Fear | Locking up and condensing the body, as well as receding into space and retracting into the shape of the body. |
Sadness | Passive weight sadness, sinking (letting the ribcage fall), head down, drooping shoulders and arm(s) to upper body, loss of muscle tone |
Joy | Jumping and rhythmic movements. Lightness (light) and free flow. Movements that enlarge the body in a horizontal and vertical direction and upward movements in space. |
Emotions and movement qualities (adapted from [30]).
These motor patterns for each emotion delimit the possible movements of the individual, determining specific dynamics of action in its structural coupling with the environment, which in turn determine the individual’s modes of relationship. Therefore, in the observation of an individual’s mode of movement it is possible to characterize these modes of relationship from the flow of postural movement dynamics generated by each emotion. These dynamics are distinguished in the experience from the proprioceptive perception, because the quality of the movements in which they generate the dynamics of activity of the muscular synergies, − speed and direction of the movement, force and muscular tone- are specific in each emotional disposition, for such in the lived experience patterns of emotional perceptions are evoked registered from proprioceptive perceptions that are correlated with the states of the evoked body.
Thus, the human being with his capacity of reflection, can recognize an emotional state in himself through the proprioceptive perception of the sensation-movement of the body of his emotion, in a present. From the study of the emotional experience, it was shown how the proprioceptive perception plays a central role in the identification of the sensations associated with global states of the body, giving emergence to the emotional experience [31]. The execution of specific body movements evokes emotional states related to those movements [32]. In turn, an emotional state modulates afferent muscular activity, modifying the patterns of sensation-movement. These observations confirm that the continuous modulation of the behavior and experience of a human being is constituted in a joint and disjointed operation of the three operational domains: body, relation and language.
In the study of Shafir et al. [30] they show that the repetition of a movement is capable of evoking an emotion, the attention is directed to the execution of that movement or sequences of movements, therefore, proprioceptive perception is active. In this way, if from the reflective movement of the attention, proprioceptive perception is intended in a present, the emotion is modulated in relation to the immediate environment and not to the flow of evocative associations of a past or future, generating a greater congruence in the structural coupling with the environment in which the living body exists, in a present.
The aforementioned is confirmed by the results of our studies about emotional plasticity in people who practice the cognitive body integration method (CBI), which correspond to a movement-based contemplative practice [33]. CBI practice is constituted from the model of the three-dimensionality of behavior to which we have made reference in this chapter. In the research we measured the autonomic response, through the pupil diameter, during the presentation of images with emotional content in a group of people who had experience in CBI practices and in a control group (CG). Our results showed that the CBI group presented shorter pupil recovery times than the CG group, showing a better emotional adaptation given the context of the individual, in a present [34].
The concept of emotional plasticity alludes to the natural loss of generative behavioral plasticity in the epigenesis of the individual, due to the history of structural links with others and the environment in which they are placed. This generates ways of moving, doing, and interpreting that are proper to the way of life of the family and culture in which the person lives, configuring in their behaviors modes of emotion that maintain a prevalence of a basic emotion, which over time restricts the domains of action of people, often reaching states of distress and loss of wellbeing within the current way of life. Thus, from the model we call “three-dimensionality of behavior”, correlations between the three operational domains of behavior are distinguished, generating correlations between ways of doing, relating and interpreting of people; from which personalized practices are designed. These practices consist of exercises in which the movement of attention towards the body – in a recursive and frequently manner- is synchronized with dynamic recurrent and recursive movements that involve the master muscles of the muscular synergies of an emotion, with reflections of what occurs in the present. Thus, these practices are intended primarily to restore emotional plasticity in people, and generate learning to modulate their emotional states, from intentional attention to proprioceptive perception, which facilitates placing oneself in the space within the environment in which one exists, maintaining a state of presence in the here and now of the body, which gives an emotional autonomy that modulates the physiological states congruent with the present contingencies, maintaining well-being in the sense of coherence with the present situation and not only of joy or enjoyment.
The purposes of approaching the paradigm from which the reflexive logic of the explanations of our observations of the phenomena of human behavior and experience is generated are, on the one hand, to show how the explanatory models and their concepts configure the perceptual objects of the world that we perceive, in this case from the doing of science. And on the other hand, to show how the recognition of the autonomy and self-reliance of the body, which reveals the knowledge that results in the continuous structural coupling of the organism with its environment, gives us a look at how the harmonies or orders that are given in the co-evolutionary drift of living species are generated, which allows us to have new references to evaluate the incidences in the individual and collective well-being of the ways of doing and relating of people in the current way of life.
In relation to our study, we can conclude that, in the epigenesis of an individual, a structural congruence is generated -between proprioception as an operation of the body and the configuration of proprioceptive perception in the domain of language- generating a co-determination of both phenomena in the structural coupling of the individual with others and his immediate environment, in a present. This explains that proprioceptive perception is not a dual phenomenon, but emerges from the interaction of the three operational domains of behavior as a coherent and unified experience. Proprioceptive perception - as the perceptual object of the observer in language - modulates and in turn is modulated by the muscular physiology that from its structural changes specifies qualities of movement observed in individual behavior and that in its experience are configured as qualities of movement, volume, relative dispositions of parts of the body and relative to their situation in space.
Proprioceptive perception has great implications for the modulation of an individual’s mode of emotion, which are defined by specific physiological states. This occurs because the dynamics of specific movements of each base emotion - which characterizes the way of moving -, are related to the conservation in adaptation of the individual within his changing environment, in a present, and not to the interpretations that he makes of his situation, which is the case of secondary or social emotions, those that do not present defined physiological and cognitive states and respond to cultural learning. Therefore, proprioceptive perception places the individual in a situation within his present circumstances, which occurs in conjunction with reflexive attentional movements of a generative character of an incorporation in the field of attention of proprioceptive perception. This attentional movement is correlated with changes in the motor surfaces that modulate their way of moving in congruence with the circumstances of the environment. This explains why contemplative practices that intend attentional reflexive movements together with body movements decrease the states of stress, which from our perspective is a generative physiological alteration of the secondary emotions that respond to the associative flow of language.
Consequently, assuming that the cognitive processes of both language and the body maintain an operational closure, we postulate that proprioceptive perception as a perceptual object is configured by spatial and movement qualities that correlate in the body domain with structural changes of the sensory and motor surfaces of the corporeal self in its interaction with the environment. Thus, the self in its history of structural coupling with its environment generates the sensorimotor learnings constitutive of the proprioceptive structure and networks of attentional selectivity that make possible the perception of a delimited internal space that originates its external space, which correspond to a space in which its existence is constituted and in which it exists, bringing to the hand the possibility of taking a perspective of itself, which occurs when differentiating proprioceptively from others and from the changing environment in which they exist. In other words, behavior and the experience of the lived world are co-determined in the interactive operation of the three operational domains of a disjointed character of behavior. And as we see the environment in which an observer distinguishes an individual, it does not correspond to his or her lived world.
From this proposal, new interesting topics are opened to deepen the understanding of these phenomena: the relations that are constituted between the reflexive movement of attention and body movements of the individual in relation to the configuration of the proprioceptive perception.
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\\n\\nThe Corresponding Author also warrants and represents that: (i) they have the full power to enter into this Publication Agreement on their own behalf and on behalf of each Co-Author; and (ii) they have the necessary rights and/or title in and to the Chapter to grant IntechOpen, on behalf of themselves and any Co-Author, the rights and licenses expressed to be granted in this Publication Agreement. If the Chapter was prepared jointly by the Corresponding Author and any Co-Author, the Corresponding Author warrants and represents that: (i) each Co-Author agrees to the submission, license and publication of the Chapter on the terms of this Publication Agreement; and (ii) they have the authority to enter into this Publication Agreement on behalf of and bind each Co-Author. The Corresponding Author shall: (i) ensure each Co-Author complies with all relevant provisions of this Publication Agreement, including those relating to confidentiality, performance and standards, as if a party to this Publication Agreement; and (ii) remain primarily liable for all acts and/or omissions of each such Co-Author.
\\n\\nThe Corresponding Author agrees to indemnify and hold IntechOpen harmless against all liabilities, costs, expenses, damages and losses and all reasonable legal costs and expenses suffered or incurred by IntechOpen arising out of or in connection with any breach of the aforementioned representations and warranties. This indemnity shall not cover IntechOpen to the extent that a claim under it results from IntechOpen's negligence or willful misconduct.
\\n\\n4.2 Nothing in this Publication Agreement shall have the effect of excluding or limiting any liability for death or personal injury caused by negligence or any other liability that cannot be excluded or limited by applicable law.
\\n\\n5. TERMINATION
\\n\\n5.1 IntechOpen has a right to terminate this Publication Agreement for quality, program, technical or other reasons with immediate effect, including without limitation (i) if the Corresponding Author or any Co-Author commits a material breach of this Publication Agreement; (ii) if the Corresponding Author or any Co-Author (being an individual) is the subject of a bankruptcy petition, application or order; or (iii) if the Corresponding Author or any Co-Author (being a company) commences negotiations with all or any class of its creditors with a view to rescheduling any of its debts, or makes a proposal for or enters into any compromise or arrangement with any of its creditors.
\\n\\nIn case of termination, IntechOpen will notify the Corresponding Author, in writing, of the decision.
\\n\\n6. INTECHOPEN’S DUTIES AND RIGHTS
\\n\\n6.1 Unless prevented from doing so by events outside its reasonable control, IntechOpen, in its discretion, agrees to publish the Chapter attributing it to the Corresponding Author and any Co-Author.
\\n\\n6.2 IntechOpen has the right to use the Corresponding Author’s and any Co-Author’s names and likeness in connection with scientific dissemination, retrieval, archiving, web hosting and promotion and marketing of the Chapter and has the right to contact the Corresponding Author and any Co-Author until the Chapter is publicly available on any platform owned and/or operated by IntechOpen.
\\n\\n6.3 IntechOpen is granted the authority to enforce the rights from this Publication Agreement, on behalf of the Corresponding Author and any Co-Author, against third parties (for example in cases of plagiarism or copyright infringements). In respect of any such infringement or suspected infringement of the copyright in the Chapter, IntechOpen shall have absolute discretion in addressing any such infringement which is likely to affect IntechOpen's rights under this Publication Agreement, including issuing and conducting proceedings against the suspected infringer.
\\n\\n7. MISCELLANEOUS
\\n\\n7.1 Further Assurance: The Corresponding Author shall and will ensure that any relevant third party (including any Co-Author) shall, execute and deliver whatever further documents or deeds and perform such acts as IntechOpen reasonably requires from time to time for the purpose of giving IntechOpen the full benefit of the provisions of this Publication Agreement.
\\n\\n7.2 Third Party Rights: A person who is not a party to this Publication Agreement may not enforce any of its provisions under the Contracts (Rights of Third Parties) Act 1999.
\\n\\n7.3 Entire Agreement: This Publication Agreement constitutes the entire agreement between the parties in relation to its subject matter. It replaces and extinguishes all prior agreements, draft agreements, arrangements, collateral warranties, collateral contracts, statements, assurances, representations and undertakings of any nature made by or on behalf of the parties, whether oral or written, in relation to that subject matter. Each party acknowledges that in entering into this Publication Agreement it has not relied upon any oral or written statements, collateral or other warranties, assurances, representations or undertakings which were made by or on behalf of the other party in relation to the subject matter of this Publication Agreement at any time before its signature (together "Pre-Contractual Statements"), other than those which are set out in this Publication Agreement. Each party hereby waives all rights and remedies which might otherwise be available to it in relation to such Pre-Contractual Statements. Nothing in this clause shall exclude or restrict the liability of either party arising out of its pre-contract fraudulent misrepresentation or fraudulent concealment.
\\n\\n7.4 Waiver: No failure or delay by a party to exercise any right or remedy provided under this Publication Agreement or by law shall constitute a waiver of that or any other right or remedy, nor shall it preclude or restrict the further exercise of that or any other right or remedy. No single or partial exercise of such right or remedy shall preclude or restrict the further exercise of that or any other right or remedy.
\\n\\n7.5 Variation: No variation of this Publication Agreement shall be effective unless it is in writing and signed by the parties (or their duly authorized representatives).
\\n\\n7.6 Severance: If any provision or part-provision of this Publication Agreement is or becomes invalid, illegal or unenforceable, it shall be deemed modified to the minimum extent necessary to make it valid, legal and enforceable. If such modification is not possible, the relevant provision or part-provision shall be deemed deleted.
\\n\\nAny modification to or deletion of a provision or part-provision under this clause shall not affect the validity and enforceability of the rest of this Publication Agreement.
\\n\\n7.7 No partnership: Nothing in this Publication Agreement is intended to, or shall be deemed to, establish or create any partnership or joint venture or the relationship of principal and agent or employer and employee between IntechOpen and the Corresponding Author or any Co-Author, nor authorize any party to make or enter into any commitments for or on behalf of any other party.
\\n\\n7.8 Governing law: This Publication Agreement and any dispute or claim (including non-contractual disputes or claims) arising out of or in connection with it or its subject matter or formation shall be governed by and construed in accordance with the law of England and Wales. The parties submit to the exclusive jurisdiction of the English courts to settle any dispute or claim arising out of or in connection with this Publication Agreement (including any non-contractual disputes or claims).
\\n\\nLast updated: 2020-11-27
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The Corresponding Author (acting on behalf of all Authors) and INTECHOPEN LIMITED, incorporated and registered in England and Wales with company number 11086078 and a registered office at 5 Princes Gate Court, London, United Kingdom, SW7 2QJ conclude the following Agreement regarding the publication of a Book Chapter:
\n\n1. DEFINITIONS
\n\nCorresponding Author: The Author of the Chapter who serves as a Signatory to this Agreement. The Corresponding Author acts on behalf of any other Co-Author.
\n\nCo-Author: All other Authors of the Chapter besides the Corresponding Author.
\n\nIntechOpen: IntechOpen Ltd., the Publisher of the Book.
\n\nBook: The publication as a collection of chapters compiled by IntechOpen including the Chapter. Chapter: The original literary work created by Corresponding Author and any Co-Author that is the subject of this Agreement.
\n\n2. CORRESPONDING AUTHOR'S GRANT OF RIGHTS
\n\n2.1 Subject to the following Article, the Corresponding Author grants and shall ensure that each Co-Author grants, to IntechOpen, during the full term of copyright and any extensions or renewals of that term the following:
\n\nThe aforementioned licenses shall survive the expiry or termination of this Agreement for any reason.
\n\n2.2 The Corresponding Author (on their own behalf and on behalf of any Co-Author) reserves the following rights to the Chapter but agrees not to exercise them in such a way as to adversely affect IntechOpen's ability to utilize the full benefit of this Publication Agreement: (i) reprographic rights worldwide, other than those which subsist in the typographical arrangement of the Chapter as published by IntechOpen; and (ii) public lending rights arising under the Public Lending Right Act 1979, as amended from time to time, and any similar rights arising in any part of the world.
\n\nThe Corresponding Author confirms that they (and any Co-Author) are and will remain a member of any applicable licensing and collecting society and any successor to that body responsible for administering royalties for the reprographic reproduction of copyright works.
\n\nSubject to the license granted above, copyright in the Chapter and all versions of it created during IntechOpen's editing process (including the published version) is retained by the Corresponding Author and any Co-Author.
\n\nSubject to the license granted above, the Corresponding Author and any Co-Author retains patent, trademark and other intellectual property rights to the Chapter.
\n\n2.3 All rights granted to IntechOpen in this Article are assignable, sublicensable or otherwise transferrable to third parties without the Corresponding Author's or any Co-Author’s specific approval.
\n\n2.4 The Corresponding Author (on their own behalf and on behalf of each Co-Author) will not assert any rights under the Copyright, Designs and Patents Act 1988 to object to derogatory treatment of the Chapter as a consequence of IntechOpen's changes to the Chapter arising from translation of it, corrections and edits for house style, removal of problematic material and other reasonable edits.
\n\n3. CORRESPONDING AUTHOR'S DUTIES
\n\n3.1 When distributing or re-publishing the Chapter, the Corresponding Author agrees to credit the Book in which the Chapter has been published as the source of first publication, as well as IntechOpen. The Corresponding Author warrants that each Co-Author will also credit the Book in which the Chapter has been published as the source of first publication, as well as IntechOpen, when they are distributing or re-publishing the Chapter.
\n\n3.2 When submitting the Chapter, the Corresponding Author agrees to:
\n\nThe Corresponding Author will be held responsible for the payment of the Open Access Publishing Fees.
\n\nAll payments shall be due 30 days from the date of the issued invoice. The Corresponding Author or the payer on the Corresponding Author's and Co-Authors' behalf will bear all banking and similar charges incurred.
\n\n3.3 The Corresponding Author shall obtain in writing all consents necessary for the reproduction of any material in which a third-party right exists, including quotations, photographs and illustrations, in all editions of the Chapter worldwide for the full term of the above licenses, and shall provide to IntechOpen upon request the original copies of such consents for inspection (at IntechOpen's option) or photocopies of such consents.
\n\nThe Corresponding Author shall obtain written informed consent for publication from people who might recognize themselves or be identified by others (e.g. from case reports or photographs).
\n\n3.4 The Corresponding Author and any Co-Author shall respect confidentiality rights during and after the termination of this Agreement. The information contained in all correspondence and documents as part of the publishing activity between IntechOpen and the Corresponding Author and any Co-Author are confidential and are intended only for the recipient. The contents may not be disclosed publicly and are not intended for unauthorized use or distribution. Any use, disclosure, copying, or distribution is prohibited and may be unlawful.
\n\n4. CORRESPONDING AUTHOR'S WARRANTY
\n\n4.1 The Corresponding Author represents and warrants that the Chapter does not and will not breach any applicable law or the rights of any third party and, specifically, that the Chapter contains no matter that is defamatory or that infringes any literary or proprietary rights, intellectual property rights, or any rights of privacy. The Corresponding Author warrants and represents that: (i) the Chapter is the original work of themselves and any Co-Author and is not copied wholly or substantially from any other work or material or any other source; (ii) the Chapter has not been formally published in any other peer-reviewed journal or in a book or edited collection, and is not under consideration for any such publication; (iii) they themselves and any Co-Author are qualifying persons under section 154 of the Copyright, Designs and Patents Act 1988; (iv) they themselves and any Co-Author have not assigned and will not during the term of this Publication Agreement purport to assign any of the rights granted to IntechOpen under this Publication Agreement; and (v) the rights granted by this Publication Agreement are free from any security interest, option, mortgage, charge or lien.
\n\nThe Corresponding Author also warrants and represents that: (i) they have the full power to enter into this Publication Agreement on their own behalf and on behalf of each Co-Author; and (ii) they have the necessary rights and/or title in and to the Chapter to grant IntechOpen, on behalf of themselves and any Co-Author, the rights and licenses expressed to be granted in this Publication Agreement. If the Chapter was prepared jointly by the Corresponding Author and any Co-Author, the Corresponding Author warrants and represents that: (i) each Co-Author agrees to the submission, license and publication of the Chapter on the terms of this Publication Agreement; and (ii) they have the authority to enter into this Publication Agreement on behalf of and bind each Co-Author. The Corresponding Author shall: (i) ensure each Co-Author complies with all relevant provisions of this Publication Agreement, including those relating to confidentiality, performance and standards, as if a party to this Publication Agreement; and (ii) remain primarily liable for all acts and/or omissions of each such Co-Author.
\n\nThe Corresponding Author agrees to indemnify and hold IntechOpen harmless against all liabilities, costs, expenses, damages and losses and all reasonable legal costs and expenses suffered or incurred by IntechOpen arising out of or in connection with any breach of the aforementioned representations and warranties. This indemnity shall not cover IntechOpen to the extent that a claim under it results from IntechOpen's negligence or willful misconduct.
\n\n4.2 Nothing in this Publication Agreement shall have the effect of excluding or limiting any liability for death or personal injury caused by negligence or any other liability that cannot be excluded or limited by applicable law.
\n\n5. TERMINATION
\n\n5.1 IntechOpen has a right to terminate this Publication Agreement for quality, program, technical or other reasons with immediate effect, including without limitation (i) if the Corresponding Author or any Co-Author commits a material breach of this Publication Agreement; (ii) if the Corresponding Author or any Co-Author (being an individual) is the subject of a bankruptcy petition, application or order; or (iii) if the Corresponding Author or any Co-Author (being a company) commences negotiations with all or any class of its creditors with a view to rescheduling any of its debts, or makes a proposal for or enters into any compromise or arrangement with any of its creditors.
\n\nIn case of termination, IntechOpen will notify the Corresponding Author, in writing, of the decision.
\n\n6. INTECHOPEN’S DUTIES AND RIGHTS
\n\n6.1 Unless prevented from doing so by events outside its reasonable control, IntechOpen, in its discretion, agrees to publish the Chapter attributing it to the Corresponding Author and any Co-Author.
\n\n6.2 IntechOpen has the right to use the Corresponding Author’s and any Co-Author’s names and likeness in connection with scientific dissemination, retrieval, archiving, web hosting and promotion and marketing of the Chapter and has the right to contact the Corresponding Author and any Co-Author until the Chapter is publicly available on any platform owned and/or operated by IntechOpen.
\n\n6.3 IntechOpen is granted the authority to enforce the rights from this Publication Agreement, on behalf of the Corresponding Author and any Co-Author, against third parties (for example in cases of plagiarism or copyright infringements). In respect of any such infringement or suspected infringement of the copyright in the Chapter, IntechOpen shall have absolute discretion in addressing any such infringement which is likely to affect IntechOpen's rights under this Publication Agreement, including issuing and conducting proceedings against the suspected infringer.
\n\n7. MISCELLANEOUS
\n\n7.1 Further Assurance: The Corresponding Author shall and will ensure that any relevant third party (including any Co-Author) shall, execute and deliver whatever further documents or deeds and perform such acts as IntechOpen reasonably requires from time to time for the purpose of giving IntechOpen the full benefit of the provisions of this Publication Agreement.
\n\n7.2 Third Party Rights: A person who is not a party to this Publication Agreement may not enforce any of its provisions under the Contracts (Rights of Third Parties) Act 1999.
\n\n7.3 Entire Agreement: This Publication Agreement constitutes the entire agreement between the parties in relation to its subject matter. It replaces and extinguishes all prior agreements, draft agreements, arrangements, collateral warranties, collateral contracts, statements, assurances, representations and undertakings of any nature made by or on behalf of the parties, whether oral or written, in relation to that subject matter. Each party acknowledges that in entering into this Publication Agreement it has not relied upon any oral or written statements, collateral or other warranties, assurances, representations or undertakings which were made by or on behalf of the other party in relation to the subject matter of this Publication Agreement at any time before its signature (together "Pre-Contractual Statements"), other than those which are set out in this Publication Agreement. Each party hereby waives all rights and remedies which might otherwise be available to it in relation to such Pre-Contractual Statements. Nothing in this clause shall exclude or restrict the liability of either party arising out of its pre-contract fraudulent misrepresentation or fraudulent concealment.
\n\n7.4 Waiver: No failure or delay by a party to exercise any right or remedy provided under this Publication Agreement or by law shall constitute a waiver of that or any other right or remedy, nor shall it preclude or restrict the further exercise of that or any other right or remedy. No single or partial exercise of such right or remedy shall preclude or restrict the further exercise of that or any other right or remedy.
\n\n7.5 Variation: No variation of this Publication Agreement shall be effective unless it is in writing and signed by the parties (or their duly authorized representatives).
\n\n7.6 Severance: If any provision or part-provision of this Publication Agreement is or becomes invalid, illegal or unenforceable, it shall be deemed modified to the minimum extent necessary to make it valid, legal and enforceable. If such modification is not possible, the relevant provision or part-provision shall be deemed deleted.
\n\nAny modification to or deletion of a provision or part-provision under this clause shall not affect the validity and enforceability of the rest of this Publication Agreement.
\n\n7.7 No partnership: Nothing in this Publication Agreement is intended to, or shall be deemed to, establish or create any partnership or joint venture or the relationship of principal and agent or employer and employee between IntechOpen and the Corresponding Author or any Co-Author, nor authorize any party to make or enter into any commitments for or on behalf of any other party.
\n\n7.8 Governing law: This Publication Agreement and any dispute or claim (including non-contractual disputes or claims) arising out of or in connection with it or its subject matter or formation shall be governed by and construed in accordance with the law of England and Wales. The parties submit to the exclusive jurisdiction of the English courts to settle any dispute or claim arising out of or in connection with this Publication Agreement (including any non-contractual disputes or claims).
\n\nLast updated: 2020-11-27
\n\n\n\n
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I am also a member of the team in charge for the supervision of Ph.D. students in the fields of development of silicon based planar waveguide sensor devices, study of inelastic electron tunnelling in planar tunnelling nanostructures for sensing applications and development of organotellurium(IV) compounds for semiconductor applications. I am a specialist in data analysis techniques and nanosurface structure. I have served as the editor for many books, been a member of the editorial board in science journals, have published many papers and hold many patents.",institutionString:null,institution:{name:"Sheffield Hallam University",country:{name:"United Kingdom"}}},{id:"54525",title:"Prof.",name:"Abdul Latif",middleName:null,surname:"Ahmad",slug:"abdul-latif-ahmad",fullName:"Abdul Latif Ahmad",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"20567",title:"Prof.",name:"Ado",middleName:null,surname:"Jorio",slug:"ado-jorio",fullName:"Ado Jorio",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Universidade Federal de Minas Gerais",country:{name:"Brazil"}}},{id:"47940",title:"Dr.",name:"Alberto",middleName:null,surname:"Mantovani",slug:"alberto-mantovani",fullName:"Alberto Mantovani",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"12392",title:"Mr.",name:"Alex",middleName:null,surname:"Lazinica",slug:"alex-lazinica",fullName:"Alex Lazinica",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/12392/images/7282_n.png",biography:"Alex Lazinica is the founder and CEO of IntechOpen. After obtaining a Master's degree in Mechanical Engineering, he continued his PhD studies in Robotics at the Vienna University of Technology. Here he worked as a robotic researcher with the university's Intelligent Manufacturing Systems Group as well as a guest researcher at various European universities, including the Swiss Federal Institute of Technology Lausanne (EPFL). During this time he published more than 20 scientific papers, gave presentations, served as a reviewer for major robotic journals and conferences and most importantly he co-founded and built the International Journal of Advanced Robotic Systems- world's first Open Access journal in the field of robotics. Starting this journal was a pivotal point in his career, since it was a pathway to founding IntechOpen - Open Access publisher focused on addressing academic researchers needs. Alex is a personification of IntechOpen key values being trusted, open and entrepreneurial. Today his focus is on defining the growth and development strategy for the company.",institutionString:null,institution:{name:"TU Wien",country:{name:"Austria"}}},{id:"19816",title:"Prof.",name:"Alexander",middleName:null,surname:"Kokorin",slug:"alexander-kokorin",fullName:"Alexander Kokorin",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/19816/images/1607_n.jpg",biography:"Alexander I. Kokorin: born: 1947, Moscow; DSc., PhD; Principal Research Fellow (Research Professor) of Department of Kinetics and Catalysis, N. Semenov Institute of Chemical Physics, Russian Academy of Sciences, Moscow.\r\nArea of research interests: physical chemistry of complex-organized molecular and nanosized systems, including polymer-metal complexes; the surface of doped oxide semiconductors. He is an expert in structural, absorptive, catalytic and photocatalytic properties, in structural organization and dynamic features of ionic liquids, in magnetic interactions between paramagnetic centers. The author or co-author of 3 books, over 200 articles and reviews in scientific journals and books. He is an actual member of the International EPR/ESR Society, European Society on Quantum Solar Energy Conversion, Moscow House of Scientists, of the Board of Moscow Physical Society.",institutionString:null,institution:{name:"Semenov Institute of Chemical Physics",country:{name:"Russia"}}},{id:"62389",title:"PhD.",name:"Ali Demir",middleName:null,surname:"Sezer",slug:"ali-demir-sezer",fullName:"Ali Demir Sezer",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/62389/images/3413_n.jpg",biography:"Dr. Ali Demir Sezer has a Ph.D. from Pharmaceutical Biotechnology at the Faculty of Pharmacy, University of Marmara (Turkey). He is the member of many Pharmaceutical Associations and acts as a reviewer of scientific journals and European projects under different research areas such as: drug delivery systems, nanotechnology and pharmaceutical biotechnology. Dr. Sezer is the author of many scientific publications in peer-reviewed journals and poster communications. Focus of his research activity is drug delivery, physico-chemical characterization and biological evaluation of biopolymers micro and nanoparticles as modified drug delivery system, and colloidal drug carriers (liposomes, nanoparticles etc.).",institutionString:null,institution:{name:"Marmara University",country:{name:"Turkey"}}},{id:"61051",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:null},{id:"100762",title:"Prof.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"St David's Medical Center",country:{name:"United States of America"}}},{id:"107416",title:"Dr.",name:"Andrea",middleName:null,surname:"Natale",slug:"andrea-natale",fullName:"Andrea Natale",position:null,profilePictureURL:"//cdnintech.com/web/frontend/www/assets/author.svg",biography:null,institutionString:null,institution:{name:"Texas Cardiac Arrhythmia",country:{name:"United States of America"}}},{id:"64434",title:"Dr.",name:"Angkoon",middleName:null,surname:"Phinyomark",slug:"angkoon-phinyomark",fullName:"Angkoon Phinyomark",position:null,profilePictureURL:"https://mts.intechopen.com/storage/users/64434/images/2619_n.jpg",biography:"My name is Angkoon Phinyomark. I received a B.Eng. degree in Computer Engineering with First Class Honors in 2008 from Prince of Songkla University, Songkhla, Thailand, where I received a Ph.D. degree in Electrical Engineering. My research interests are primarily in the area of biomedical signal processing and classification notably EMG (electromyography signal), EOG (electrooculography signal), and EEG (electroencephalography signal), image analysis notably breast cancer analysis and optical coherence tomography, and rehabilitation engineering. I became a student member of IEEE in 2008. During October 2011-March 2012, I had worked at School of Computer Science and Electronic Engineering, University of Essex, Colchester, Essex, United Kingdom. In addition, during a B.Eng. I had been a visiting research student at Faculty of Computer Science, University of Murcia, Murcia, Spain for three months.\n\nI have published over 40 papers during 5 years in refereed journals, books, and conference proceedings in the areas of electro-physiological signals processing and classification, notably EMG and EOG signals, fractal analysis, wavelet analysis, texture analysis, feature extraction and machine learning algorithms, and assistive and rehabilitative devices. I have several computer programming language certificates, i.e. Sun Certified Programmer for the Java 2 Platform 1.4 (SCJP), Microsoft Certified Professional Developer, Web Developer (MCPD), Microsoft Certified Technology Specialist, .NET Framework 2.0 Web (MCTS). 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