eng. coste ioan dumitru study of protein and oil … · 2011. 10. 21. · eng. coste ioan dumitru...
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UNIVERSITY OF AGRICULTURAL
SCIENCE AND VETERINARY MEDICINE
CLUJ-NAPOCA
DOCTORAL SCHOOL
FACULTY OF AGRICULTURE
Eng. COSTE IOAN DUMITRU
STUDY OF PROTEIN AND OIL CONTENT VARIABILITY IN A SERIES OF
LOCAL AND SYNTHETIC POPULATIONS, INBRED
LINES AND HYBRIDS OF MAIZE
(SUMMARY OF THE PhD THESIS)
SCIENTIFIC ADVISOR
PROF. DR. IOAN HAŞ
CLUJ-NAPOCA
2011
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TABLE OF CONTENTS INTRODUCTION ...................................................................................................................... 9-5 CHAPTER I.............................................................................................................................. 11-7 IMPORTANCE OF MAIZE.................................................................................................... 11-7
1.1. ECONOMIC IMPORTANCE .......................................................................................... 11-7 1.2. WORLDWIDE SPREAD................................................................................................. 13-8 1.3. WOLDWIDE MAIZE YIELD.......................................................................................... 14-8 1.4. MAIZE CULTIVATED AREAS AND YIELD IN ROMANIA......................................... 16-8 1.5. MAIZE UTILIZATION ...................................................................................................... 19 1.6. SPECIFIC UTILIZATION OF PROTEIN AND OIL FROM MAIZE .................................. 21
CHAPTER II ................................................................................................................................22 GENETICS AND IMPROVEMENT OF OIL CONTENT ........................................................22
2.1. INHERITANCE OF OIL CONCENTRATION IN MAIZE.................................................. 22 2.2. ASSOCIATION OF OIL CONTENT WITH OTHER AGRONOMIC TRAITS ................... 23 2.3. HIGH-OIL MAIZE BREEDING PROGRAMS ................................................................... 24 2.4. ENVIRONMENTAL EFFECTS ON MAIZE OIL CONTENT ............................................ 26 2.5. XENIA EFFECTS ON OIL CONTENT............................................................................... 27 2.6. SELECTION PROGRESS FOR INCREASED OIL CONTENT IN MAIZE ........................ 28 2.7. SELECTION METHODS USED TO INCREASE OIL CONTENT IN MAZIE................... 29
2.7.1. Phenotipic recurent selections for increased oil content ............................................... 29 2.7.2. Single kernel selection for increased oil content. .......................................................... 30 2.7.3. Intensive selection for increased oil content ................................................................. 31 2.7.4. Modified phenotipic recurent selection for increased oil content .................................. 32
2.8. FACTORS THAT CONTROL OIL QUALITY IN MAIZE HYBRIDS................................ 33 2.8.1. Factors that affect oilty acid composition of maize oil .................................................. 36 2.8.2. Inheritance of fatty acid compositon in maize ............................................................... 38
2.9. VITAMIN E OR TOCOPHEROL ....................................................................................... 42 2.9.1. Genetic variability........................................................................................................ 42
CHAPTER III...............................................................................................................................44 GENETICS AND IMPROVEMENT OF PROTEIN QUALITY AND CONTENT IN MAIZE
.......................................................................................................................................................44 3.1. BIOCHEMICAL CHARACTERIZATION AND GENETIC BASE OF PROTEIN CONTENT IN MAIZE ................................................................................................................................. 45
3.1.1. Mutant genes for protein quality................................................................................... 45 3.1.2. Inheritance of protein quality and content .................................................................... 47 3.1.3. Pleiotropic effects of mutant genes for protein quality .................................................. 48
3.2. GERMPLASM FOR PROTEIN QUALITY AND CONTENT IMPROVEMENT................ 49 3.3. DEVELOPMENT OF MAIZE TYPES RICH IN SUPERIOR QUALITY PROTEINS......... 52
3.3.1. Problemes of opaque-2 maize....................................................................................... 55 3.3.2. Breeding maize types with hard endosperm rich in essential aminoacids...................... 59
3.4. STRUCTURE AND IMPORTANCE OF ZEIN FROM MAIZE PROTEIN ......................... 66 3.4.1. Descovery of protein bodies ......................................................................................... 66
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3.4.2. Origin of prolaminic and globulinic protein bodies ...................................................... 68
CHAPTER IV........................................................................................................................... 71-9 BIOLOGIC MATERIAL AND RESEARCH METHODS.................................................... 71-9
4.1. BIOLOGIC MATERIAL ................................................................................................. 71-9 4.2. PREPEARING THE MATERIAL FOR CHEMICAL ANALYSIS ...................................... 72 4.3. NIR INSTALAB 600 ANALIZER....................................................................................... 73 4.4. NATURAL CONDITIONS OF THE EXPERIMENTAL SITE............................................ 75 4.5. STATISTICAL ANALYSES OF THE RESULTS............................................................... 80 4.6. OBSERVATIONS, NOTES AND DETEMINATIONS....................................................... 82 4.7. OBJECTIVES................................................................................................................ 83-10
CHAPTER V .......................................................................................................................... 84-12 RESULTS AND DISCUSSIONS ( I ) ................................................................................... 84-12
5.1. PROTEIN AND OIL CONTENT IN THE COLLECTION OF LOCAL POPULATIONS OF MAIZE FROM ARDS TURDA ............................................................................................ 84-12 5.2. PROTEIN AND OIL CONTENT IN SYNTHETIC POPULATIONS OF MAIZE CREATED AT ARDS TURDA.................................................................................................................... 87 5.3. PROTEIN AND OIL CONTENT IN INBRED LINES CREATED AT ARDS TURDA .. 90-12 5.4. STUDY OF THE RELATIONS BETWEEN KERNEL CONSTITUENTS IN LOCAL POPULATIONS........................................................................................................................ 97 5.5. STUDY OF THE RELATIONS BETWEEN KERNEL CONSTITUENTS IN SYNTHETIC POPULATIONS.......................................................................................................................102 5.6. STUDY OF THE RELATIONS BETWEEN KERNEL CONSTITUENTS IN INBRED LINES........................................................................................................................................... 107-13 5.7. STUDY OF PROTEIN AND OIL CONTENT IN THE COLECTION OF INBRED LINES FROM ARDS TURDA DEPENDING ON KERNEL TYPE......................................................112
5.7.1. Protein and oil conten in inbred lines depending on kernel type ..................................112 5.7.2. Study of the relations between kernel constituents depending on kernel type................119
CHAPTER VI....................................................................................................................... 138-19 RESULTS AND DISCUSSIONS ( II ) ................................................................................. 138-19
6.1. PROTEIN CONTENT IN ISONUCLEAR INBRED LINES IN A BALANCED GENETIC SYSTEM............................................................................................................................ 138-19 6.2. OIL CONTENT IN ISONUCLEAR INBRED LINES IN A BALANCED GENETIC SYSTEM............................................................................................................................ 139-20 6.3. PROTEIN CONTENT IN ISONUCLEAR INBRED LINES IN A UNBALANCED GENETIC SYSTEM(2009-2010)......................................................................................................... 142-22 6.4. OIL CONTENT IN ISONUCLEAR INBRED LINES IN A UNBALANCED GENETIC SYSTEM(2009-2010)......................................................................................................... 145-23
CHAPTER VII ..................................................................................................................... 151-25 INHERITANCE OF AGRONOMIC AND QUALITY TRAITS OF KERNELS IN CERTAIN
ISONUCLEAR INBRED LINE CROSSES ........................................................................ 151-25 7.1. GENETIC STUDY ON DETERMINISM OF EAR WEIGHT.............................................152
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7.2. GENETIC STUDY ON DETERMINISM OF EAR LENGTH ...................................... 157-26 7.3. GENETIC STUDY ON DETERMINISM OF THE NUMBER OF ROWS PER EAR..........161 7.4. GENETIC STUDY ON DETERMINISM OF 1000 KERNELS WEIGHT...........................167 7.5. GENETIC STUDY ON DETERMINISM OF YIELD CAPACITY.....................................173 7.6. GENETIC STUDY ON DETERMINISM OF DRY MATTER CONTENT IN KERNELS..179 7.7. GENETIC STUDY ON THE FALLING RESISTANCE OF MAIZE STALKS...................185 7.8. GENETIC STUDY ON DETERMINISM OF PROTEIN AND OIL CONTENT IN SELF POLLINATED AND OPEN POLLINATED KERNELS .................................................... 190-28
7.8.1. Variance of protein content ................................................................................... 190-28 7.8.2. Study on protein content inheritance ..................................................................... 193-30 7.8.3. Variance of oil content .......................................................................................... 205-32 7.8.4. Study on oil content inheritance ............................................................................ 209-33
CHAPTER VIII.................................................................................................................... 222-35 STUDY ON PROTEIN AND OIL CONTENT INHERITANCE IN A DIALLEL SYSTEM
TYPE P(P-1) ......................................................................................................................... 222-35 8.1. PROTEIN CONTENT IN SELF POLLINATED KERNELS ........................................ 223-36 8.2. PROTEIN CONTENT IN OPEN POLLINATED KERNELS .............................................227 8.3. OIL CONTENT IN SELF POLLINATED KERNELS ................................................. 230-40 8.4. OIL CONTENT IN OPEN POLLINATED KERNELS.......................................................233
CHAPTER IX....................................................................................................................... 237-43 CONCLUSIONS AND RECOMMENDATIONS ............................................................... 237-43 BIBLIOGRAPHY ................................................................................................................ 243-46
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INTRODUCTION
Maize is one of the most important crops due to its high productivity and its multiple
uses as a food source for humans, livestock feed and as raw material in various industries
(VASILICĂ, 1991, quoted by MUNTEAN et all., 2008).
Maize kernels are used in industry to obtain alcohol, starch, dextrin, glucose as well
as other products like plastics, glue, acetone, dyes, etc. From the seed embryo a good quality
dietetic oil can be extracted that prevents the accumulation of cholesterol in the blood
(MUNTEAN et all., 2008)
The study of the maize germoplasm from ARDS Turda was initiated in an effort to
identify local and synthetic populations and inbred lines which have potential use in the
improvement of protein and oil content in maize kernel. There were also studied a series of
isonuclear lines to identify if the utilization of different types of cytoplasm affect protein and
oil content.
Experimentation was carried out for a period of two years (2009-2010) in the
experimental field of the corn breeding laboratory from Turda, and the testing of isonuclear
lines regarding yield capacity, dry matter content and falling resistance has been made in
two years (2009-2010) and two locations. Some of the expenses were sustained from a PN
II-52-129/2008 project, developed in collaboration of UASVM Cluj-Napoca and ARDS
Turda.
The present thesis is structured in nine chapters and contains 254 pages, 103 tables,
76 figures, 24 conclusions, 5 recommendations and 155 bibliographical titles were quoted
from the scientific literature.
The completion of this thesis, witch concludes a milestone in my professional
training, would not have been possible without the help of some dear and wonderful people.
Using this occasion I would like to thank them.
I wish to express my gratitude to the scientific advisor, Mr. Prof. Dr. Ioan Haş, for his
patience, guidance, professional training and constant attention during the years of my PhD
preparation.
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I would also like to thank Mrs. Dr. eng. Voichiţa Haş, head of the Corn Breeding
Laboratory from ARDS Turda for her support given in order to carry out my experiments. I
thank the entire staff of the Corn Breeding Laboratory from ARDS Turda, who contributed
to my professional training and gave me their support to carry out my experiments
I thank my colleagues from the Department of Genetics and Plant Breeding, for their
friendship and support given in these three years spent together.
Special thanks and gratitude to my family, I want to thank my wife for here patience
and understanding, moral support given throughout my PhD training.
Special thanks to the leadership of the University of Agricultural Sciences and
Veterinary Medicine, Doctoral School, who contributed to my professional training.
.
Thank you!
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CHAPTER I
IMPORTANCE OF MAIZE
1.1. ECONOMIC IMPORTANCE
On a worldwide scale maize is situated on the third place concerning cultivated area
(MUNTEAN et all., 2001; CRISTEA, 2004) and in first place concerning yield
capacity(TROYER, 2006). Maize is also a valuable raw material for industry: from its kernels
there can be extracted oil, starch, alcohol, glucose and other products such as syrup, pectin,
dextrin, plastics, lactic acid, acetic acid, acetone, dyes, synthetic rubber etc. Maize stalk can be
used to produce paper, cardboard, nitrocellulose (CRISTEA, 2004; ŞTEFAN, 2004).
About one fifth of world maize production is used directly for human consumption.
The tendency to reduce direct consumption of maize in human diet is more obvious in
developed countries, where its ratio represents only 6.8%, while in developing countries is
60% (CRISTEA, 2004).
In addition to traditional uses, the overproduction of maize in the U.S. has led to its
use in the production of bioethanol, maize thus becoming one of the plants that will
contribute to saving fossil fuels and maintaining a cleaner environment (HAŞ, 2004).
The world's corn production is used mostly for animal feed (72%) and current trends
indicate that this share will increase. Regarding this manner of use we can observe the
differences between developed and developing countries. The developed countries use about
88% of the maize production for animal fodder, in the developing countries it represents
only 27.9%. In these countries the situation is better in terms of maize use in food-
processing industry. Currently, about 5% of world production of maize is used in industrial
processing (CRISTEA, 2004).
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1.2. WORLDWIDE SPREAD
Corn is grown in various climatic and soil conditions, in the northern hemisphere in
Canada and Russia up to 58 ° latitude, and in the southern hemisphere, in New Zealand, up
to 42-43 degrees. Grown for grain, corn can be found between 42º and 53º northern latitude,
this limit is exceeded when the destination use is as green fodder.
1.3. WORLDWIDE MAIZE YIELD
Due to more efficient application of technology and the use of heterosis phenomenon
in creating new maize hybrids, in order to increase corn production its yield, in some
countries reached a spectacular level (CRISTEA, 2004).
Estimated by FAO, in 2004 total maize kernel production was about 724.589.004
tons, and average global yield per hectare to 49.2 q / ha (FAO, 2004). The largest areas
planted with maize were in America, followed by Asia, Africa and Europe.
1.4. MAIZE CULTIVATED AREAS AND YIELD IN ROMANIA
Regarding the evolution of maize production in Romania before 1960 the average yield
per area unit was somewhere around 1000 kg. In this period different varieties and local
populations were used. The introduction of double-cross hybrids, after 1960 provides a
doubling of maize production
After the Revolution in December 1989 due to new situations created by structural and
functional reorganization of state institutions, and the transition to market economy,
agriculture has undergone major transformations too, which unfortunately, didn’t have a
favorable influence on agricultural yield, in all situations.
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CHAPTER II
BIOLOGIC MATERIAL AND RESEARCH METHODS
2.1. BIOLOGIC MATERIAL A total of 422 inbred lines, 258 of local populations and 60 synthetic populations of
maize were studied in terms of protein, oil, starch and fiber content. This collection of
germplasm from ARDS Turda consists of material from its own breeding program and
materials obtained from biological material exchange with other research centers.
In addition to the 422 inbred lines, 258 local populations and 60 synthetic
populations, five groups of isonucleare lines have been studied (in each group an elite inbred
line and six sister lines on different types of cytoplasm) and were crossed with three testers
for TC221, TB367, D105 group and four testers for the group TC209 and TC243 to make an
experimental systems that enables the testing of the role of cytoplasm, testers and testers x
cytoplasm interactions.
The studied isonucleare lines (seven in each group) were obtained by transferring the
nucleus on different types of cytoplasm using 10 backcrosses (Figure 1). It was considered
that in this way the transfer was made to about 99.9%, the only difference between the
isonuclear lines from the same group consisted in genetic factors situated in the cytoplasm
and eventual interactions between the nuclear and cytoplasm factors.
The isonuclear lines were created between 1992-2004 at ARDS Turda, starting from
the assumption that between different sources of cytoplasm there could be differences
regarding the genetic value. After this period the isonuclear lines were maintained by self
pollination or using SIB pollination (CHICINAŞ et all., 2009).
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Cytoplasm Nucleus donor donor
I X 50.00 % II X 75.00 % III X
87.50 % IV X 93.75 % V X
96.88 % VI X 98.44 % VII X
99.22 % VIII X 99.61 % IX X
99.90 % X
Fig. 1. Transferring method of the nucleus on to a new cytoplasm
2.3. OBJECTIVES The objectives were as follows:
study of the local population, synthetic population and inbred lines of maize from
ARDS Turda regarding protein and oil content, and identification of correlations
between the kernel constituents
identification of the protein and oil rich/poor genotypes from the germplasm
collection;
study of the general and specific combining ability of the isonuclear inbred lines in
crosses with elite tester inbred lines;
to evidence the influence of cytoplasm on maize ear traits, protein and oil content,
yield capacity, falling resistance and dry matter content;
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identification of valuable types of cytoplasm that can be used in breeding programs to
improve oil and protein content;
to highlight the cytoplasm effects involved in the inheritance of protein and oil
content using direct and reciprocal crosses.
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CHAPTER III
RESULTS AND DISCUTIONS (I)
3.1. PROTEIN AND OIL CONTENT IN THE COLLECTION OF LOCAL POPULATIONS OF MAIZE FROM ARDS TURDA
Statistical analysis of protein and oil content is presented in table 1. Mean protein
content in local populations was 13.69% and oil content was 5.38%.
Range for protein was 4.40% and 3.50% for oil content. Confidence level for the
local populations was 0.10% for protein content and 0.08% for oil content.
Table 1
Statistical analysis of protein and oil content for a series of local maize populations Statistical parameters Protein Oil
Mean % 13.69 5.38 Standard error 0.05 0.04 Standard deviation 0.84 0.63 Sample variance 0.70 0.39 Range 4.40 3.50 Minimum value 11.20 3.80 Maximum value 15.60 7.30 The coefficient of variation 6.12 11.6 Confidence level 95% 0.10 0.08 Number of cases 258 258
3.2. PROTEIN AND OIL CONTENT IN INBRED LINES CREATED AT ARDS TURDA Statistical analysis of protein and oil content of the 422 inbred lines created at ARDS
Turda is presented in table 2.
Mean protein content is 13.43%, the range for protein content being 6.90%. Variance
was relatively low (1.17%) and standard deviation was 1.08%, resulting that for a large
number of inbred lines the deviation from the average was statistically significant.
Mean oil content was 4.16%, sample variance was 0.75% and confidence level was
0.08%. Oil content range has been between 2.40% and 7.60%.
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Table 2
Statistical analysis of protein and oil content for a series of inbred maize lines Statistical parameters Protein Oil
Mean % 13.43 4.16 Standard error 0.05 0.04 Standard deviation 1.08 0.87 Sample variance 1.17 0.75 Range 6.90 5.20 Minimum value 10.80 2.40 Maximum value 17.70 7.60 The coefficient of variation 8.06 20.88 Confidence level 95% 0.10 0.08 Number of cases 422 422
3.3. STUDY OF THE RELATIONS BETWEEN KERNEL CONSTITUENTS IN INBRED LINES
Figure 2 presents the relationship between protein and oil content for the 422 inbred
lines from the germplasm collection of maize from ARDS Turda. The correlation coefficient
between protein content and oil content (r = 0.3866 **) indicate a distinctly significant
positive relationship between the two characters. The relationship is described by the upward
slope of the regression line b = 0.4816, analyzing the regression it can be seen that by
increasing the oil content with a unit protein content increases by 0.48 units.
The coefficient of determination (R2 = 0.1495) shows that in the case of the 422
inbred lines of maize studied, the variation of protein content is determind in a share of 14%
by the variation in oil content. Analyzing the distribution of inbred lines, two inbred lines
with high protein and oil content were distingwished (TC106 with 16.4% protein and 7.5%
oil, T442 15.6% protein and 7.2% oil) giving them potential use in breeding programs to
improve protein and oil content.
The correlation between protein and starch content (r =- 0.6263 **) shown in figure 3
is dinsctincly significantlty negative and the coefficient of detemination (R2 = 0.3922) shows
that the variation of protein content is determined in share of 39% by starch content
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variation. Following the downward slope of the regression line (b = -0.2507) it can be seen
that increasing the starch content with a uint causes a decrease in protein content with 0.25
units. From the analyzed inbred lines regarding the correlation between protein and starch
content three inbred lines were highlighted because of high protein and starch content T397
(16% protein and 65.4% starch), TC350 (15.7% proteins and 66.6% starch) and TC245
(15.7% protein and 66.3% starch).
In case of the correlation between protein and fiber content shown in figure 4 (r =
0.6748 **) there can be identified a distinctly significant positive correlation, with an
upward slope of the regression line (b = 0.7931), and it can be that increasing the fiber
content with a unit protein content is increased with 0.79 units. Analyzing the relation
between protein and fiber content it can be concluded that inbred lines T139 (17.7% protein
and 7.1% fiber), TC106 (16.4% protein and 7.1% fiber) and TB370 (16% 6.9 protein and
fiber) can be used in maize breeding programs for high protein and fiber content.
0
2
4
6
8
10
12
14
16
18
20
2 3 4 5 6 7 8
oil(%)
port
ein(
%)
r = 0.3866**
R2= 0.1495
y = 11.4215+0.4816x
n = 422
Fig. 2. Relation between protein and oil content in inbred maize lines
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0
2
4
6
8
10
12
14
16
18
20
40 45 50 55 60 65 70 75
starch(%)
prot
ein(
%)
r = -0.6263**
R2= 0.3922y = 30.3619-0.2507x
n = 422
Fig. 3. Relation between protein and starch content in inbred maize lines
0
2
4
6
8
10
12
14
16
18
20
2 3 4 5 6 7 8
fiber(%)
port
ein(
%)
r = 0.6748**
R2= 0.4553
y = 9.5555+0.7931x
n = 422
Fig. 4. Relation between protein and fiber content in inbred maize lines
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0
1
2
3
4
5
6
7
8
7 9 11 13 15 17 19
protein(%)
oil(%
)
r = 0.3866**
R2= 0.1495
y = -0.006+0.3104x
n = 422
Fig. 5. Relation between oil and protein content in inbred maize lines
0
1
2
3
4
5
6
7
8
40 45 50 55 60 65 70 75
starch(%)
oil(%
)
r = -0.8454**
R2= 0.7147
y = 22.5164-0.2717x
n = 422
Fig. 6. Relation between oil and starch content in inbred maize lines
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0
1
2
3
4
5
6
7
8
2 3 4 5 6 7 8
fiber%)
oil(%
)
r = 0.6579**
R2= 0.4329
y = 1.1315+0.6208x
n = 422
Fig. 7. Relation between oil and fiber content in inbred maize lines
Figure 5 shows the relationship between oil and protein content. Correlation
coefficient value, r = 0.3866 ** indicates a distinctly significant positive relationship
between the studied characters. The relationship between these constituents of the maize
kernel is described by an upward regression slope b = 0.3104, expressing a close relationship
between oil and protein content, thus increasing protein content with a unit increases oil
content with 0.31 units. The coefficient of determination R2 = 0.1495 shows that the
variation of oil content is determined at a rate of 14% by variation in protein content. Inbred
lines that are recommended to be used in breeding programs to improve oil and protein
content are TC106 (7.5% oil and 16.4 protein) and T442 (7.2% oil and 15.6% protein).
The correlation between oil and starch content (r =- 0.8454 **) shown in figure 6 is
distinctly significantly negative and oil content variation is determined in 71% by the
variation of starch content. Following the downward slope of the regression line (b = -
0.2717) it can be seen that increasing the starch content with a unit causes a decrease in oil
content by 0.27 units. Based on analysis of regressions two inbred lines have been identified
(TC344 6.2% oil and 65.1% starch, 5.9% TC346 65.3% oil and starch) with high oil and
starch content.
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In case of the correlation between oil and fiber content shown in figure 7 (r = 0.6579
**) we can identify a distinclty significant positive correlation with an upward slope of the
regression line (b = 0.6208), and it can be observed that increasing the fiber content with a
unit oil content increases by 0.62 units. Analyzing the oil content according to fiber content
it can be observed that inbred lines TC344 (7.5% oil and 7.5% fiber), TC344A (7.6% oil and
7.2% fiber), TC106 (7, 5% oil and 7.1% fiber) and TC375 (7.1% oil and 7.3% fiber) can be
used in breeding programs to increase oil and fiber content.
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CHAPTER IV
RESULTS AND DISCUTIONS (II)
4.1. PROTEIN CONTENT IN ISONUCLEAR INBRED LINES IN A BALANCED GENETIC SYSTEM
Table 3 shows the protein content in five groups of isonuclear maize lines. The mean
protein content of the five groups of isonuclear lines was 12.86%, the highest protein content
being recorded for group D105 with cit.TB329 (14.47%) while the lowest for the group of
lines TC316 with cit.TC221 (11.53%). Out of the cytoplasm’s that were used to improve the
protein content only cit.T248 showed positive deviation from conventional cytoplasm in
four of the five groups of isonuclare lines. Using cytoplasm from TC221 the protein content
decreased with 0.29% presenting a distinctly significant negative difference.
Table 3
Protein content of five groups of isonuclear inbred maize lines in a balanced genetic system (ARDS Turda, 2010)
Nucleus Cytoplasm TC209
% TC316
% TC243
% TB367
% D105
%
Mean for each type of
cytoplasm
Deviation+/-from conventional
cytoplasm
Conventional 12.23 12.40 12.87 13.33 13.63 12.89 0.00 cit.T248 13.13*** 12.83* 13.90*** 12.13ooo 13.80 13.16 +0.27**
cit.TB329 13.07*** 11.83oo 12.93 12.13ooo 14.47*** 12.89 0.00 cit.TC177 12.67* 13.23*** 12.13ooo 12.37ooo 13.47 12.77 -0.12 cit.TC221 12.30 11.53ooo 13.60*** 13.07 12.50ooo 12.60 -0.29oo
Mean of isonuclear lines 12.68 12.37oo 13.09 12.61 13.57*** 12.86
LDS comparisons C P 5% = 0.17 P 1% = 0.23 P 0.1% = 0.30
LDS comparisons N P 5% = 0.28 P 1% = 0.41 P 0.1% = 0.62
LDS comparisons C×N P 5% = 0.39 P 1% = 0.52 P 0.1% = 0.68
From the five groups of isonucleare lines, the highest mean protein content was
registered for the lines generated by the D105. Specific interactions were noted between the
type of cytoplasm and nucleus. The inbred lines TC209 (cit. T248), TC209 (cit.TB329),
TC209 (cit.TC177), TC316 (cit.T248), TC316 (cit.TC177), TC243 (cit.T248), TC243
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(cit.T221) , D105 (cit. TB329) exceeded with statistically significant values the protein
content of lines supplying the nucleus, while for the lines TC316 (cit.TC221), TC316
(cit.TB329), TC243 (cit.TC177), TB367 (cit.T248), TB367 (cit.TB329), TB367 (cit.TC177)
and D105 (cit.TC221) protein content was significantly lower than that of lines that supplied
the nucleus.
4.2. OIL CONTENT IN ISONUCLEAR INBRED LINES IN A BALANCED GENETIC SYSTEM
Oil content of maize kernels of a system of inbred lines with five types of cytoplasm
is shown in table 4. Experimental system average was 4.92%, the highest values of oil content were recorded in the group of isonucleare lines TB367 (6.45%), followed by the
group of isonucleare lines TC243 (5.25%).
Table 4
Oil content of five groups of isonuclear inbred maize lines in a balanced genetic system (ARDS Turda, 2010)
Nucleus Cytoplasm
TC209 %
TC316 %
TC243 %
TB367 %
D105 %
Mean for each type of cytoplasm
Deviation+/- from
conventional cytoplasm
Conventional 4.00 3.90 5.37 6.07 4.73 4.81 0.00
cit.T248 4.60** 3.90 5.20 5.83 4.87 4.88 +0.07 cit.TB329 3.67 4.07 5.33 6.57* 4.70 4.87 +0.05 cit.TC177 4.37 3.80 5.43 7.30*** 4.70 5.12 +0.31** cit.TC221 4.40 4.40* 4.93o 6.47 4.40 4.92 +0.11
Mean of isonuclear lines
4.21ooo 4.01ooo 5.25* 6.45*** 4.68o 4.92
LDS comparisons C P 5% = 0.20 P 1% = 0.26 P 0.1% = 0.35
LDS comparisons N P 5% = 0.23 P 1% = 0.33 P 0.1% = 0.50
LDS comparisons C×N P 5% = 0.44 P 1% = 0.59 P 0.1% = 0.77
Cytoplasm type had a higher influence on oil content in case of the cytoplasm from
the inbred line TC177. Range of the oil content was between 3.67% for TC209 (cit. TB329)
and 7.30% for TB367 (cit.TC177).
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Favorable specific interactions between cytoplasm and nucleus were recorded for the
following inbred lines: TC209 (cit.T248), TC316 (cit.TC221), TB367 (cit.TC177) and
TB367 (cit.TB329). In one case, the transfer of the nucleus led to a significant decrease in
oil content for the inbred line TC243 (cit.TC221).
Table 5 Analysis of variance of the chemical composition of the maize kernel for five groups of
inbred lines in a balanced genetic system (ARDS Turda, 2010) Protein Oil Cause of variability DF SP s2 SP s2
Genotypes (G) 24 36.95 1.54** 64.39 2.68**
Nucleus(N) (4) 13.50 3.37** 57.45 14.36**
Cytoplasm(C) (4) 2.50 0.63** 0.84 0.21*
Interaction N x C (16) 20.94 1.31** 6.10 0.38**
Repetitions(R) 2 0.11 0.06 0.05 0.03
Error N 8 0.91 0.11 0.58 0.07
Error C 40 2.20 0.06 2.84 0.07 Total 74 40.18 67.86
As it can be observed from table 5, the genetic factors of cytoplasm have a distinctly
significant influence on variability of protein content (s2 = 0.63 **) and a significant
influence on variability of oil content (s2 = 0.21 *). Nuclear factors have a distinctly
significant influence on the variability of protein content (3.37 **) and oil content (14.36
**). The nuclear-cytoplasm interactions have a distinctly significant influence on protein
content variability (1.31 **) and significant influence on oil content variability (0.38 *).
In figure 8 the share of genetic factors involved in protein and oil content variability
is presented, the share of nuclear factors in the variability of protein content being 37% and
89% for oil content variability. Regarding the influence of cytoplasm, the share was 7% for
protein content variability and 1% for oil content variability. Nuclear-cytoplasm interactions
had a 57% share in the variability in protein content and 10% in oil content variability.
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22
1- nucleus; 2- cytoplasm; 3- nucleus-cytoplasm interaction
Fig. 8. Contribution of nucleus, cytoplasm and interactions between nucleus and cytoplasm in the variance of protein and oil content of five groups of isonuclear inbred maize lines in a
balanced genetic system (ARDS Turda, 2010)
4.3. PROTEIN CONTENT IN ISONUCLEAR INBRED LINES IN A UNBALANCED GENETIC SYSTEM (2009-2010)
The results of the analyses for protein content from self pollinated and open
pollinated kernels between 2009-2010 in the group of isonuclear lines TC243 is shown in
table 6.
Average protein content of the two experimental years, for the self pollinated kernels
was 12.63% with quite large differences between the two experimental years (13.49% in
2009 and 11.79% in 2010). In the case of open pollinated kernels the average protein content
of the two years was 11.41%, and the differences between the experiment years was lower
(11.77% in 2009 and 11.05% in 2010).
In comparison wih the control line TC243 that generated the group of isonuclear lines
with a protein content of 11.64%, five isonucleare lines registered an increase in protein
content TC243 (cit.T248), TC243 (cit.TC208), TC243 (cit.TC221), TC243 (cit.K1080) and
TC243 (cit.K2051). Analysis of differences in the two types of pollinated material provided
data that is statistically assured differences indicate that the protein rich isonucleare lines
T248, TC208, TC221, K1080 and K2051 can be used to improve the protein content of the
line TC243 that generated the group of isonuclear lines.
Protein
37%
7%57%
1 2 3
Oil
89%
1% 10%
1 2 3
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23
Table 6
Protein content of the tested isonuclear inbred maize lines TC243 in a unbalanced genetic system (ARDS Turda, 2009-2010)
Protein content of maize kernel from self pollinated ears (%)
Protein content of maize kernel from open pollinated ears (%)
Mean for each cytoplasm (%)
Isonuclear inbred line
2009
2010
Mean % vs initial
line
2009 2010 Mean % vs initial
line
Mean content
Relative
value
TC243(martor/control) 13.27 12.07 12.67 100.0 11.93 9.27 10.60 100.0 11.64 100.0 TC243(cit. A665) 13.30 11.77 12.54 98.9 10.77 9.27 10.02 94.5 11.28 96.9 TC243(cit.T248) 14.20 12.27 13.24*** 104.5 11.27 11.30 11.29*** 106.5 12.26 105.4 TC243(cit.TC208) 12.93 11.90 12.42 98.0 12.17 11.73 11.95*** 112.7 12.18 104.7 TC243(cit.TC221) 13.47 11.87 12.67 100.0 12.00 11.23 11.62*** 109.6 12.14 104.4 TC243(cit.K1080) 14.07 10.93 12.50 98.7 11.83 12.13 11.98*** 113.0 12.24 105.2 TC243(cit.K2051) 13.20 11.53 12.37 97.6 12.40 12.43 12.42*** 117.1 12.39 106.5 Mean of isonuclear lines 13.49 11.76 12.63 11.77 11.05 11.41 12.02 LDS (P=5%) 0.32 0.26 LDS (P=1%) 0.43 0.35 LDS (P=0.1%) 0.55 0.45
4.4. OIL CONTENT IN ISONUCLEAR INBRED LINES IN A UNBALANCED GENETIC SYSTEM (2009-2010)
The results of the analysis for oil content in the group of isonucleare lines TB367 are
shown in table 7. Mean oil content of this group of lines was the highest from the five
groups of isonuclear lines studied.
Compared to the control line TB367 that generated the group of isonuclear lines with
an oil content of 5.60%, three isonucleare lines registered an increase in oil content TB367
(cit.TC209), TB367 (cit.TC208) and TB367 (cit.TC221) and had the following amounts of
oil content: 6.39%, 6.25%, 6.04%. These values allow us to say that these genotypes may be
considered rich in oil content. Analysis of differences between the two types of pollination
provided statistically assured results for the isonuclear lines with high oil content (COSTE et
al., 2011).
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24
Table 7 Oil content of the tested isonuclear inbred maize lines TB367 in an unbalanced genetic
system (ARDS Turda, 2009-2010) Oil content of maize kernel from
self pollinated ears (%) Oil content of maize kernel from
open pollinated ears (%) Mean for each cytoplasm (%)
Isonuclear inbred line
2009
2010
Mean
% vs initial
line
2009
2010
Mean %
vs initial line
Mean content
Relative value
TB367 (control) 6.13 5.70 5.92 100.0 5.83 4.73 5.28 100.0 5.60 100.0 TB367(cit.T248) 5.40 6.10 5.75 97.2 5.27 5.63 5.45 103.2 5.60 100.0 TB367(cit.TB329) 5.50 5.83 5.67 95.8 5.00 7.30 6.15*** 116.5 5.91 105.5 TB367(cit.TC208) 6.17 6.60 6.39 107.9 5.07 7.17 6.12*** 115.9 6.25 111.7 TB367(cit.TC221) 6.03 5.77 5.90 99.7 5.67 6.70 6.19*** 117.1 6.04 107.9 TB367(cit.TC209) 6.40 5.97 6.19 104.6 7.93 5.27 6.60*** 125.0 6.39 114.2 TB367(cit.K2051) 6.47 6.53 6.50* 109.9 5.60 5.07 5.34 101.0 5.92 105.7 Mean of isonuclear lines 6.01 6.07 6.04 5.77 5.98 5.87 5.96 LDS (P=5%) 0.48 0.49 LDS (P=1%) 0.64 0.65 DL (P=0.1%) LDS (P=0.1%) 0.83 0.84
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25
CHAPTER V INHERITANCE OF AGRONOMIC AND QUALITY TRAITS OF KERNELS
IN CERTAIN GROUPS OF ISONUCLEAR INBRED LINE CROSSES
The literature indicates that most of the characters that help to increase yield in maize
(ear size, ear length, number of rows of kernels per ear, number of kernels per row, 1000
kernel weight) are genetically conditioned, in highest share at nuclear level, but there are
allegations that in the inheritance of some of these characters genes with cytoplasm
localization could be involved (HALLAUER and MIRANDA, 1981; CĂBULEA, 1975;
CĂBULEA et all., 1981; STUBER and EDWARDS, 1986; STUBER et all., 1992; HAŞ,
1992; CĂBULEA et all., 1994; CĂBULEA et all., 1999; TROYER, 2001; CĂBULEA,
2004; SARCA, 2004; HAŞ,2004).
The understanding of kernel quality determinism is important in combination with
improving kernel production and other agronomic characteristics, but also for improvement
of nutritional and industrial properties (POLLAK and SCOTT, 2005; OSORNO and
CARENA, 2008). Increasing the nutritional value of maize grain is possible using
technologic and genetic methods (HAŞ et all., 2004; HEGYI et all., 2007; HEGYI et
all.,2008; IDIKUT et all., 2009).
Yield capacity can be considered the most complex trait from genetic point of view.
Due to the considerable number of genes involved, and gene interactions, the heterozygous
state, as well as the complex interactions with environmental factors, make yield capacity
one of the most difficult targets as a genetic study and is one of the traits with the lowest
heritability (HALLAUER and MIRANDA, 1981).
Genetic determinism of protein and zein content is significantly correlated at an
additive genetic level and is controlled by additive gene and cytoplasm actions (CĂBULEA
and ZETEA, 1973) or only by gene interactions (CĂBULEA et al., 1984).
Given the importance of embryo size in the improvement of oil content, the oil
content is related to kernel size and hence in the inheritance of oil content cytoplasm actions
are involved (ALEXANDER and CREECH, 1977; COSMIN et al., 1993; SARCA et al.,
1995).
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26
5.1. GENETIC STUDY ON DETERMINISM OF EAR LENGTH Analysis of variance for ear length for the five groups of isonucleare maize lines that
were tested is presented in table 8. Variance for the influence of experimental years was
distinctly significant for the group of isonuclear lines TC209 and TC243 and significant for
TC221 and TB367, revealing the importance of climatic conditions in the manifestation of
this character.
Table 8 Analysis of variance of ear length for the tested isonuclear inbred maize lines
(ARDS Turda, 2009-2010) Isonuclear line
TC209 Isonuclear line
TC243 Isonuclear line
TC221 Isonuclear line
TB367 Isonuclear line
D105 Cause of variability DF
SP s2 SP s2
DF
SP s2 SP s2 SP s2
Experimental years (Y) 1 192.71 192.71** 84.86 84.86* 1 60.32 60.32* 19.06 19.06* 0.00 0.00
Genotype 27 211.51 7.83** 333.65 12.36** 20 48.19 2.41* 17.09 0.85* 30.03 1.50**
Cytoplasm (C) (6) 67.85 11.31** 4.24 0.71 (6) 13.17 2.20* 3.71 0.62 3.79 0.63
Tester (T) (3) 117.65 39.22** 305.89 101.96** (2) 17.83 8.91 5.92 2.96 14.91 7.46**
(C×T) interaction (18) 26.01 1.45** 23.52 1.31** (12) 17.19 1.43 7.46 0.62 11.33 0.94**
(Y×T) interaction 3 13.08 4.36** 8.30 2.77* 2 3.53 1.77 7.49 3.74* 2.24 1.12**
(Y×C) interaction 6 21.85 3.64** 12.31 2.05** 6 12.05 2.01 7.61 1.27* 2.18 0.36
(Y×T×C) interaction 18 23.89 1.33* 12.45 0.69 12 23.61 1.97* 8.18 0.68 4.38 0.37
Repetition ( R ) 2 5.28 2.64 5.19 2.60 2 8.45 4.23 0.21 0.11 1.93 0.97
Error Y 2 2.04 1.02 1.85 0.93 2 2.60 1.30 1.32 0.66 1.34 0.67
Error T 12 5.88 0.49 9.20 0.77 8 19.92 2.49 7.10 0.89 0.64 0.08
Error C 96 59.12 0.62 52.06 0.54 72 68.45 0.95 31.39 0.44 24.55 0.34
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27
1- the share of cytoplasm effect variance;
2 - the share tester effect variance; 3 the share "cytoplasm x tester" interaction effect variance Fig. 9. Share of each factor involved in the genetic variance of ear length for the five
groups of isonuclear inbred lines that were tested
Cytoplasm have provided statistically significant differences in two of the five groups
tested, higher values of variance were recorded for testers, and for cytoplasm × tester
interactions lower values were recorded but statistically significant.
Share of the factors involved in ear length variance is presented in figure 9.
Cytoplasm share ranged from 1% for the group of lines TC243 and 32% for the group of
lines TC209, testers share was 35% for the group of lines TB367 and 92% for the group of
lines TC243; "cytoplasm × testers" interactions had a share ranging between 7% in the group
of lines TC243 and 43% in the group of lines TB367.
The results of general and specific combining ability involved in the inheritance of
ear length for the group of isonuclear lines TC209 is presented in table 9. Range of
cytoplasm effects was between -0.69cm for TC209 and 0.82cm for TC209(cit.TC177), the
general combining ability effects due to testers ranged between -0.52cm for TB329 and 1.45
cm for Lo3Rf. Values of specific combining effects ranged between -0.9cm in the hybrid
combination TC209 (cit.W633) × TC344 and 0.7cm for TC209 (cit.W633) × Lo3Rf.
TC209
32%
56%
12%
1 2 3
TC243
1%
92%
7%
1 2 3
TC221
27%
37%
36%
1 2 3
TB367
22%
35%
43%
1 2 3
D105
13%
49%
38%
1 2 3
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28
From the hybrid combinations the highest values for ear length obtained in the following hybrid combinations:
- TC209(cit.W633)×Lo3Rf=21.3cm=μ(+18.4cm)+ĝcit(+0.73cm)+ĝtest(+1.45cm)+ŝcit
× test(0.7cm)
- TC209(cit.D105)×Lo3Rf=20.9cm=μ(+18.4cm)+ ĝcit(+0.62cm)+ ĝtest(+1.45cm)+ŝcit
× test(0.5cm)
In the inheritance of ear length for the hybrid combinations with the highest ear
length are involved in addition to the mean effects, general combining ability effects due to
testers followed by general combining ability of cytoplasm and specific combining ability.
Table 9 General and specific combining ability involved in the inheritance of the ear length in the
tested isonuclear inbred maize lines TC 209(ARDS Turda, 2009-2010) TC344 Lo3Rf TB329 TD233
cytoplasm/tester cm ŝ cit x test cm ŝ cit x test cm ŝ cit x test cm ŝ cit x test
x ĝcit
TC209 18.0 0.7 18.6 -0.6 17.5 0.3 16.8 -0.4 17.7 -0.69 TC209(cit.A665) 17.5 0.2 19.2 -0.1 17.5 0.2 17.1 -0.2 17.8 -0.61 TC209(cit.T291) 17.1 -0.4 19.2 -0.3 17.7 0.2 18.1 0.5 18.0 -0.38 TC209(cit.T248) 17.5 0.1 19.1 -0.2 17.2 -0.2 17.9 0.4 17.9 -0.49 TC209(cit.W633) 17.8 -0.9 21.3 0.7 18.7 0.1 18.8 0.1 19.1 0.73 TC209(cit.TC177) 19.4 0.6 20.7 0.0 18.5 -0.2 18.4 -0.4 19.2 0.82 TC209(cit.D105) 18.3 -0.2 20.9 0.5 18.2 -0.3 18.7 0.1 19.0 0.62
Mean for tester 17.9 19.9 17.9 18.0 18.4 0.00 ĝtest -0.48 1.45 -0.52 -0.45 18.4 0.00
LDS 5% comparison ĝcytoplasm 0.45 LDS 5% comparison ĝtester 0.33 LDS 5% comparison for interaction c × t 0.90
5.2. GENETIC STUDY ON DETERMINISM OF PROTEIN AND OIL CONTENT IN SELF POLLINATED AND OPEN POLLINATED KERNELS
5.2.1. Variance of protein content
Maize protein is less important than that of other grains due to high level of zein
fraction, but it is hoped that this inconvenience will be eliminated; still it is desirable to have
available forms with high protein content and better quality.
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29
Analysis of variance in protein content for the five groups of isonuclear lines that
were self pollinated is presented in table 10. In all five comparative trials the differences
between the two experimental years 2009 and 2010 were statistically assured.
Table 10 Analysis of variance of the protein content for the self pollinated isonuclear inbred maize
lines that were tested (ARDS Turda, 2009-2010) Isonuclear line
TC209 Isonuclear line
TC243
Isonuclear line
TC221 Isonuclear line
TB367 Isonuclear line
D105
Cause of variability DF
SP s2 SP s2
DF
SP s2 SP s2 SP s2
Experimental years (Y) 1 68.15 68.15** 21.07 21.07** 1 0.09 0.09* 8.49 8.49* 104.60 104.60** Genotype 27 39.69 1.47** 55.48 2.05** 20 31.65 1.58** 17.91 0.90** 42.65 2.13**
Cytoplasm (C) (6) 8.66 1.44** 10.92 1.82** (6) 4.76 0.79** 11.96 1.99** 9.12 1.52** Tester (T) (3) 13.83 4.61** 25.87 8.62** (2) 14.78 7.39** 2.56 1.28** 26.68 13.34**
(C×T) interaction (18) 17.21 0.96** 18.69 1.04** (12) 12.11 1.01** 3.38 0.28** 6.84 0.57** (Y×T) interaction 3 7.86 2.62** 26.32 8.77** 2 3.05 1.52** 8.90 4.45** 12.37 6.19** (Y×C) interaction 6 6.59 1.10** 2.64 0.44** 6 26.79 4.47** 7.08 1.18** 7.11 1.18**
(Y×T×C) interaction 18 11.65 0.65** 6.79 0.38** 12 15.18 1.27** 4.81 0.40** 6.45 0.54** Repetition ( R ) 2 0.06 0.03 0.16 0.08 2 0.18 0.09 0.19 0.10 0.02 0.01
Error Y 2 0.03 0.01 0.16 0.08 2 0.01 0.00 0.49 0.25 0.08 0.04 Error T 12 0.70 0.06 1.42 0.12 8 1.49 0.19 0.35 0.04 0.28 0.04 Error C 96 4.65 0.05 5.50 0.06 72 4.89 0.07 7.80 0.11 4.86 0.07
In all five test groups there were differences between genotypes and the differences
were distinctly significant statistically. Cytoplasms have provided statistically significant
differences in all five test groups, with an evident role of the cytoplasm in the differentiation
of the genotypes. The highest values were recorded for testers variance; lower values, but
distinctly significant statistically were shown by the „nucleus-cytoplasm” interactions.
In all five experimental situations interaction with experimental years was statistically
significant. Share of the factors involved in the variance of protein content is shown in figure
10. Influence of cytoplasm on the variance of protein content was between 15% and 22%.
As expected, the share was higher for the testers, between 14% for TB367 and 63% for
D105, quite high values were also recorded in most situations for cytoplasm × nucleus interactions, with values between 16% for D105 and 43% for TC209 (COSTE et al., 2011).
-
30
1- the share of cytoplasm effect variance;
2 - the share tester effect variance; 3 - the share „cytoplasm × tester” interaction effect variance Fig. 10. Share of each factor involved in the genetic variance of protein contet for the
five groups of self pollinated isonuclear inbred lines that were tested 5.2.2. Study on protein content inheritance
General combining ability effects for the cytoplasm and testers and the specific
combining capacity due to „cytoplasm x testers” interaction for the group of lines TC243 is
shown in table 11.
Average protein content in the tested isonuclear inbred lines generated by TC243
inbred line was 11.6%. With statistically significantly higher average values of content, thus
with higher values of transmission at cytoplasm level stood the isonuclear lines: TC243
(cit.TC221) - ĝcit = 0.29% and TC243-ĝcit =0.25%; in both cases the difference from ĝcit
registered for control is statistically significant. With negative values for the transmission of
protein content were noted TC243 (cit.A665)-ĝcit =- 0.40% and TC243 (cit.TC208) - ĝcit =
-0.34%.
Among the inbred tester lines, highest combining capacity at an additive level was
registered for the tester line from indurata convariety Lo3Rf with an overall combining
TC209
43%22%
35%
1 2 3
TC243
34%20%
47%
1 2 3
TC221
38%
15%
47%
1 2 3
TB367
19%
14%67%
1 2 3
D105
16% 21%
63%
1 2 3
-
31
capacity of 0.55% and the lowest value was registered for the tester inbred line TC344 -
0.56%
For the hybrid combinations, the values of specific combining ability ranged from -
0.71% for the hybrid combination TC243 (cit.A665) × TC344 to 0.83% for the hybrid
combination TC243 × TC344. The results of this comparative trial showed that the share of
cytoplasm effects, additive effects of the testers and interactions of „cytoplasm × tester”
would be relatively close, but that the importance of cytoplasm’s and testers would be
decisive in determinism of the protein content of hybrids:
- TC243(cit.K1080)×Lo3Rf= 12.6% = μ(+11.6%)+ ĝcit(+0.18%)+ ĝtest(+0.55%)+ŝcit x
test(0.26%)
- TC243(cit.TC221)×Lo3Rf=12.5%=μ(+11.6%)+ĝcit(+0.29%)+ĝtest(+0.55%)+ŝcit × test (0.10%)
Table 11
General and specific combining ability involved in the inheritance of protein content for the self pollinated isonuclear inbred maize lines that were tested
TC243 (ARDS Turda, 2009-2010) TC344 Lo3Rf TB329 TD233
cytoplasm/tester % ŝ cit ×test % ŝ cit ×test % ŝ cit ×test % ŝ cit ×test
x ĝcit
TC243 11.9 0.83 12.2 0.02 11.1 -0.51 11.3 -0.34 11.7 0.06 TC243(cit.A665) 9.9 -0.71 12.0 0.26 11.6 0.40 11.3 0.04 11.2 -0.40 TC243(cit.T248) 11.4 0.43 11.8 -0.33 11.3 -0.30 11.8 0.20 11.6 -0.03
TC243(cit.TC208) 10.6 -0.13 11.8 -0.01 11.3 0.09 11.3 0.06 11.2 -0.34 TC243(cit.TC221) 11.0 -0.36 12.5 0.10 12.0 0.13 12.0 0.13 11.9 0.29 TC243(cit.K1080) 11.1 -0.14 12.6 0.26 12.1 0.34 11.3 -0.47 11.8 0.18 TC243(cit.K2051) 11.4 0.08 12.1 -0.31 11.7 -0.15 12.2 0.38 11.8 0.25
Mean for tester 11.0 12.1 11.6 11.6 11.6 0.00 ĝtest -0.56 0.55 -0.01 0.02 11.6 0.00
LDS 5% comparison ĝcytoplasm 0.14 LDS 5% comparison ĝtester 0.16 LDS 5% comparison for interaction c × t 0.27
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32
5.3.3. Variance of oil content
Oil content in maize is important; but when the kernels are used in the industry to
manufacture alcohol, bioethanol and fodder, corn oil and soluble fiber are a byproduct of
major importance that must be exploited.
Analysis of variance in oil content of the five groups of self pollinated isonuclear
lines is presented in table 12. From this table one can conclude that the experimental years
didn’t provide statistically significant values except for the TC221 isonuclear group, the
other studied test groups showed no influence of the experimental years on the oil content.
Table 12 Analysis of variance of the oil content for the self pollinated isonuclear inbred maize lines
that were tested (ARDS Turda, 2009-2010) Isonuclear line TC209 Isonuclear line TC243 Isonuclear line TC221 Isonuclear line TB367 Isonuclear line D105 Cause of variability DF
SP s2 SP s2
DF
SP s2 SP s2 SP s2
Experimental years (Y) 1 0.05 0.05 0.00 0.00 1 9.45 9.45** 0.52 0.52 0.46 0.46
Genotype 27 27.75 1.03** 40.36 1.49** 20 6.16 0.31** 5.54 0.28 6.08 0.30**
Cytoplasm (C) (6) 1.95 0.33** 0.27 0.04 (6) 1.88 0.31** 3.20 0.53** 0.81 0.14**
Tester (T) (3) 24.39 8.13** 38.63 12.88** (2) 3.52 1.76** 0.88 0.44* 3.46 1.73**
(C×T) interaction (18) 1.41 0.08* 1.47 0.08* (12) 0.75 0.06 1.46 0.12 1.81 0.15
(Y×T) interaction 3 0.45 0.15 0.96 0.32** 2 0.76 0.38 0.55 0.28 2.44 1.22**
(Y×C) interaction 6 1.75 0.29** 1.13 0.19** 6 4.23 0.70** 0.74 0.12 2.28 0.38**
(Y×T×C) interaction 18 1.83 0.10** 3.34 0.19** 12 1.04 0.09 2.49 0.21* 1.73 0.14
Repetition ( R ) 2 0.36 0.18 0.01 0.00 2 0.47 0.23 0.39 0.19 0.22 0.11
Error Y 2 0.09 0.04 0.04 0.02 2 0.07 0.04 0.27 0.14 0.38 0.19
Error T 12 1.27 0.11 0.38 0.03 8 1.15 0.14 1.00 0.13 0.25 0.03
Error C 96 4.37 0.05 4.47 0.05 72 7.38 0.10 7.24 0.10 7.52 0.10
Cytoplasms have provided statistically significant differences in four of the five test
groups, with an important role in the differentiation of genotypes. The highest values of
variance were recorded for testers being distinctly significant in all five groups of
isonucleare lines. Statistically significant values were also recorded for the interactions
between cytoplasm × experimental years, years × testers and interaction of the three factors
(years × testers × cytoplasm).
-
33
The share of factors involved in the inheritance of oil content for the self pollinated
ears is presented in figure 11 (model of LEIN quoted by CEAPOIU, 1969).
Cytoplasm’s influence on the variance of oil content was between 1% and 58%. As
expected the share for the testers was higher and ranged from 16% for the group of TB367
isonuclear lines and 96% for the group of TC243 isonuclear lines. Relativly high values of
cytoplasm × nucleus interaction was recorded for the group of TC243 isonuclear lines (4%)
and 30% for the group of D105 isonuclear lines.
1- the share of cytoplasm effect variance;
2 - the share tester effect variance; 3 - the share "cytoplasm × tester" interaction effect variance Fig. 11. Share of each factor involved in the genetic variance of oil content for the
five groups of self pollinated isonuclear inbred lines that were tested
5.4.4. Study of oil content inheritance
The effects of general and specific combining ability involved in the inheritance of oil
content for the group of isonuclear lines generated by TB367 (self pollinated) are presented
in table 13.
TC209
5% 7%
88%
1 2 3
TC243
4%1%
96%
1 2 3
TC221
12%31%
57%
1 2 3
TB367
26%
16%58%
1 2 3
D105
30%13%
57%
1 2 3
-
34
Average oil content of the experimental system was 5%. The effects of general
combining ability at cytoplasm level ranged from -0.32% for TB367 (cit.TB329) and 0.22%
for TB367 (cit.TC208).
Among the inbred tester lines, the highest combining ability was provided by the
TC209 inbred line (0.12%) and the lowest was recorded for T291 line (-0.07%).
For the hybrid combinations, values of the specific combining ability ranged from -
0.19% for the TB367 (cit.TC221) × TD233 hybrid combination to 0.23% for the TB367
(cit.TC209) × TD233 hybrid combination.
In the determination of oil content in hybrids with the highest values have contributed
primarily the mean effects, but also equally important were the cytoplasm effects, nucleus
and „nucleus-cytoplasm” interactions.
- TB367(cit.TC208)×TC209=5.4%=μ(5%)+ĝcit(0.22%)+ĝtest(0.12%)+ŝcit×test (0.04%)
- TB367(cit.TC221)×TC209=5.3%=μ(5%)+ĝcit(0.11%)+ĝtest(0.12%)+ŝcit×test (0.08%)
Table 13 General and specific combining ability involved in the inheritance of oil content for the self
pollinated isonuclear inbred maize lines that were tested TB367 (ARDS Turda, 2009-2010)
T291 TC209 TD233 cytoplasm/tester
% ŝ cit ×test % ŝ cit ×test % ŝ cit ×test x ĝcit
TB367 4.8 -0.05 5.3 0.18 4.8 -0.13 5.0 -0.08 TB367(cit.T248) 4.9 0.02 5.1 -0.02 5.0 0.01 5.0 -0.04
TB367(cit.TB329) 4.8 0.12 4.8 -0.07 4.6 -0.04 4.7 -0.32 TB367(cit.TC208) 5.1 -0.05 5.4 0.04 5.2 0.02 5.2 0.22 TB367(cit.TC221) 5.2 0.16 5.3 0.08 4.9 -0.19 5.1 0.11 TB367(cit.TC209) 5.0 -0.07 5.1 -0.16 5.3 0.23 5.1 0.06 TB367(cit.K2051) 4.9 -0.09 5.2 -0.04 5.2 0.12 5.1 0.06
Media testeri Mean for tester 5.0 5.1 5.0 5.0 0.00
ĝtest -0.07 0.12 -0.04 5.0 0.00 LDS 5% comparison ĝcytoplasm 0.21 LDS 5% comparison ĝtester 0.18 LDS 5% comparison for interaction c × t 0.36
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CAPITOLUL VI STUDY OF PROTEIN AND OIL CONTENT INHERITANCE IN A DIALLEL
SYSTEM TYPE P (P-1) Maize proteins are not of a very high quality due to a large content of zein, a protein
fraction insoluble in water, soluble in alcohol. However the improvement of maize protein
quality is of great interest for all breeding programs, latest research efforts aiming to
improve the ratio of protein fractions, by other techniques than using opaque-2 and floury-2
genes (SARCA, 2004).
Increasing protein content was the subject of research works for nearly a century.
ALEXANDER (1988) presents that after reviewing the Illinois program, initiated by
Hopkins at the end of XIX century, the variability for this character is not exhausted, after
76 generations of selection, the frequency of favorable alleles increased by 37%. Negative
correlations were found between production capacity and protein content, but CĂBULEA et
al. (1984) argues that the two traits can be improved simultaneously.
The literature shows that in the genetic determinism of oil content variance of the
general combining ability (GCA) was higher than specific combining ability (SCA),
indicating a preponderance of additive genetic variance. Therefore to improve the oil content
different methods of selection are used, most frequently, the recurrent selection (MISEVIC
and ALEXANDER, 1989; MISOVIC et al. 1990, HAŞ et al., 2004).
Oil content was analyzed both from self pollinated ear samples and from open
pollinated ear samples. Differences between the two origins of the analyzed kernels should
not be so great, because after ALEXANDER (1988) about 83% of the oil content of the
maize kernel comes from the embryo.
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6.1. PROTEIN CONTENT IN SELF POLLINATED KERNELS
Gene effects involved in the protein content determinism for kernels from self
pollinated hybrids in a diallel system, values are presented in tables 14-16.
The average protein content for the two experimental years was 11.89%, with values
between 11.11% for TC209 × TC344 hybrid and 13% for hybrid T291 × K1080. Statistically
significant values that exceeded the average of the experimental system were reached by:
T248 x T291 (0.50%), T248 × TC209 (0.38%), T291 × TC344 (0.34%) hybrids.
Between the mean of the two experimental years there were statistically significant
differences, 12.71% in 2009 and 11.07 in 2010. Maximum protein content of 13.40% was
recorded fore T248xT291 hybrid in 2009 and the minimum value of 9.99% for the
TC209xTC344 hybrid in 2010.
Between direct and reciprocal hybrids there were statistically significant differences,
the highest values were recorded in the following hybrids:
- T248×TC209, lower protein content of 0.87% for the direct hybrid
compared to the reciprocal hybrid;
- TC209×TC344, lower protein content of 0.70% for the direct hybrid
compared to the reciprocal hybrid;
- TC209×K1080, lower protein content of 0.76% for the direct hybrid
compared to the reciprocal hybrid;
- TC208×T209, higher protein content of 0.75% compared to the reciprocal
hybrid;
- T291×TC344, higher protein content of 0.47% compared to the reciprocal
hybrid.
Differences between direct and reciprocal hybrids could result from the reciprocal
effects of the „nucleus-cytoplasm” interaction (values between -0.57 and +0.43) and to a
lesser extent, maternal effects, which for some parental forms have values that must be taken
into account (K1080-m = 0.08; T291-m = 0.05; TC344-m =- 0.07; TC209-m =- 0.05).
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37
The highest general combining ability for transmission of high protein content has
been registered for the T291 inbred line 12, 21% (ĝi = +0.32).
Low values for the transmission of protein content (additive gene effects with minus
sign) occurred in TC208 inbred lines 11. 62% (ĝi =- 0.28) and TC209 11. 66% (ĝi =- 0, 24).
Unadditive effects for some hybrid combinations had high absolute values,
highlighting the importance of specific combining capacity in the determinism (high or low)
of protein content
- T248xTC209-ŝij=+0.52;
- T291xK1080- ŝij=+0.49;
- TC208xK1080- ŝij=+0.33;
- TC209xTC344- ŝij=-0.35;
- T291xTC209- ŝij=-0.35;
- T248xK1080- ŝij=-0.27;
We can conclude that in the protein content determinism of the self pollinated maize
ears additive gene effects, unadditivee gene effects and the reciprocal effects of the
„nucleus-cytoplasm” interactions are of equal importance, the maternal gene effects being
less important.
Concerning the protein level in the determination of the optimal hybrid combination
formula it is necessary to test both direct hybrid and the reciprocal one.
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38
Table 14 Protein content of the direct and reciprocal hybrids from a diallel system p (p-1) and the
genetic effect rij of the specific interactions between the maternal cytoplasm and the nuclear genes (ARDS Turda, 2009-2010, self pollinated)
% Protein content
(%)
crt.nr. Single cross
hybrid A × B B × A Difference rij 2009 2010 Mean
content
% compared
to the system mean medie mean
1 T248 × T291 12.62 12.17 0.45 0.31 13.40 11.38 12.39 104.2 2 T248 × T208 11.62 11.42 0.20 -0.08 12.29 10.75 11.52 96.9 3 T248 × TC209 11.83 12.70 -0.87 -0.57 12.72 11.82 12.27 103.2 4 T248 × TC344 11.88 11.52 0.36 0.07 12.82 10.59 11.70 98.4 5 T248 × K1080 11.97 12.05 -0.08 0.02 13.22 10.80 12.01 101.0 6 T291 × TC208 11.87 11.72 0.15 -0.24 12.49 11.10 11.79 99.2 7 T291 × TC209 11.73 11.52 0.21 -0.17 12.15 11.10 11.63 97.8 8 T291 × TC344 12.47 12.00 0.47 -0.01 12.67 11.80 12.23 102.9 9 T291 × K1080 13.05 12.95 0.10 -0.02 13.25 12.75 13.00 109.3 10 TC208 × TC209 11.80 11.05 0.75 0.43 12.30 10.55 11.43 96.1 11 TC208× TC344 10.98 11.68 -0.70 -0.27 12.34 10.34 11.34 95.3 12 TC208 × K1080 12.08 11.93 0.15 0.33 12.89 11.14 12.01 101.0 13 TC209 × TC344 11.27 10.95 0.32 0.24 12.23 9.99 11.11 93.4 14 TC209 × K1080 11.47 12.23 -0.76 -0.13 12.73 10.97 11.85 99.6 15 TC344 × K1080 11.98 12.22 -0.24 0.12 13.15 11.05 12.10 101.8 Mean 11.91 11.87 0.03 12.71 11.07 11.89 100.0 LDS 5% comp. years 0.06 LDS 5% comp. hybrids 0.28 LDS comp. int years × hybrids 0.39
Comparison of rij to rji - 0.0492
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Table 15 Additive genetic effects (ĝi) and the maternal effects involved in the inheritance of protein content from a diallel crossing system p (p-1) (ARDS Turda, 2009-2010, self pollinated)
The constant inbred line ♀ % ĝi (m)
♂ % ĝj(t)
CCG % ĝ Maternal effects
T248 11.98 0.09 11.97 0.08 11.98 0.09 0.01 T291 12.26 0.37 12.16 0.27 12.21 0.32 0.05
TC208 11.60 -0.29 11.63 -0.26 11.62 -0.28 -0.01 TC209 11.60 -0.29 11.71 -0.18 11.66 -0.24 -0.05 TC344 11.63 -0.26 11.76 -0.13 11.70 -0.20 -0.07 K1080 12.28 0.39 12.11 0.22 12.19 0.30 0.08
System mean 11.89 11.89 11.89 Comparison of ĝi to ĝj - 0.01467 Comparison of mi to mj - 0.00984
Table 16 Mean protein content and the unadditive gene effects in a diallel system p (p-1)
(ARDS Turda, 2009-2010, self pollinated)
T 248 T 291 TC 208 TC 209 TC 344 K 1080 T 248 12.62 11.62 11.83 11.88 11.97
ŝij 0.10 -0.18 0.52 -0.08 -0.27 r 0.31 -0.08 -0.57 0.07 0.02
T 291 12.17 11.87 11.73 12.47 13.05 ŝij -0.14 -0.35 0.22 0.49 r -0.24 -0.17 -0.01 -0.02
TC 208 11.42 11.72 11.8 10.98 12.08 ŝij 0.05 -0.09 0.09 r 0.43 -0.27 0.33
TC 209 12.70 11.52 11.05 11.27 11.47 ŝij -0.35 -0.11 r 0.24 -0.13
TC 344 11.52 12.00 11.68 10.95 11.98 ŝij 0.10 r 0.12
K 1080 12.05 12.95 11.93 12.23 12.22 ŝij r
Comparison of ŝij to ŝik - 0.04428 Comparison of ŝij to ŝkl - 0.02952
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6.2. OIL CONTENT IN SELF POLLINATED KERNELS
Oil content and gene effects involved in the determinism of this character for the self
pollinated kernels are presented in tables 17-19.
Average content for the hybrids obtained in the diallel system was 4.65%, with no
differences between the two experimental years. The hybrid with the highest oil content was
T248xTC344 (5.01%), followed by TC208xTC344 (4.99%) the lowest content was recorded
in the hybrid combination T291xK1080 (4.09%). Between direct and reciprocal hybrids has
been an average difference of 0.09%; but the difference between direct and reciprocal
individual hybrids presented higher amplitudes:
- T291xTC208 difference between direct and reciprocal hybrid was -0,58%;
- TC208xTC209 difference between direct and reciprocal hybrid was -0,33%;
- T248xTC209 difference between direct and reciprocal hybrid was -0,30%;
Differences between direct and reciprocal hybrids are mostly determined by „nucleus-
cytoplasm” effects with values between -0.25 for the T248xTC209 hybrid combination and
+0.23 for the T291xTC344 hybrid combination. The comparisons between unadditive gene
effects and reciprocal effects of the interaction of nuclear genes and cytoplasm are presented
in table 19, it can be seen that in many cases, reciprocal effects were higher than the
unadditive gene effects.
In the determinism of phenotypic differences for the amount of oil between reciprocal
and direct hybrids maternal gene effects are also involved, but with lower value then the
reciprocal effects.
General combining ability effects are presented in table 18. Highest oil content is
transmitted by the TC344 inbred line (x = 4.92%, ĝi = 0.27). The lowest oil content is
transmitted by the T291 inbred lines (x = 4.50%, ĝi =- 0.15) and K1080 (x = 4.69%, ĝi =-
0.14).
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41
From the presented data we conclude that in heredity of oil content (in kernels from
the self pollinated ears) additive gene effects, reciprocal effects of „nucleus-cytoplasm”
interactions and unaditive gene effects are involved.
Table 17 Oil content of the direct and reciprocal hybrids from a diallel system p (p-1) and the genetic efect rij of the specific interactions between the maternal cytoplasm and the nuclear genes
(ARDS Turda, 2009-2010, self pollinated)
% Oli content
(%)
crt.nr. Single cross
hybrid A × B B × A Difference rij 2009 2010 Mean
content
% compared
to the system mean medie mean
1 T248 × T291 4.60 4.83 -0.23 -0.22 4.82 4.62 4.72 101.5 2 T248 × T208 4.72 4.78 -0.06 -0.09 4.80 4.70 4.75 102.2 3 T248 × TC209 4.52 4.82 -0.30 -0.25 4.87 4.47 4.67 100.4 4 T248 × TC344 5.10 4.92 0.18 0.19 5.07 4.95 5.01 107.8 5 T248 × K1080 4.67 4.63 0.04 -0.08 4.75 4.55 4.65 100.1 6 T291 × TC208 4.25 4.83 -0.58 -0.22 4.58 4.50 4.54 97.7 7 T291 × TC209 4.33 4.35 -0.02 0.01 4.22 4.47 4.34 93.4 8 T291 × TC344 4.82 4.80 0.02 0.23 4.88 4.74 4.81 103.4 9 T291× K1080 4.15 4.03 0.12 0.08 4.04 4.15 4.09 88.1 10 TC208 × TC209 4.35 4.68 -0.33 -0.23 4.45 4.58 4.52 97.1 11 TC208 × TC344 4.90 5.08 -0.18 0.04 5.00 4.99 4.99 107.4 12 TC208 × K1080 4.40 4.32 0.08 -0.03 4.24 4.49 4.36 93.8 13 TC209 × TC344 4.83 4.85 -0.02 0.14 4.73 4.95 4.84 104.1 14 TC209 × K1080 4.40 4.57 -0.17 -0.13 4.32 4.65 4.48 96.4 15 TC344 × K1080 4.98 4.92 0.06 -0.17 4.85 5.05 4.95 106.5 Mean 4.60 4.69 -0.09 4.64 4.66 4.65 100.0 LDS 5% comp. years 0.19 LDS 5% comp. hybrids 0.24 LDS comp. int years × hybrids 0.33
Comparison of rij to rji - 0.038208
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Table 18 Additive genetic effects (ĝi) and the maternal effects involved in the inheritance of oil
content from a diallel crossing system p (p-1) (ARDS Turda, 2009-2010, self pollinated)
The constant inbred line ♀ % ĝi (m)
♂ % ĝj(t)
CCG % ĝ Maternal effects
T248 4.72 0.07 4.80 0.15 4.76 0.11 -0.04 T291 4.48 -0.17 4.52 -0.13 4.50 -0.15 -0.02
TC208 4.65 0.00 4.61 -0.04 4.63 -0.02 0.02 TC209 4.62 -0.03 4.52 -0.12 4.57 -0.08 0.05 TC344 4.93 0.28 4.91 0.27 4.92 0.27 0.01 K1080 4.49 -0.15 4.88 -0.13 4.69 -0.14 -0.01
System mean 4.65 4.71 4.68 Comparison of ĝi to ĝj - 0.011463 Comparison of mi to mj - 0.007642
Table 19 Mean oil content and the unadditive gene effects in a diallel system p (p-1)
(ARDS Turda, 2009-2010, self pollinated)
T 248 T 291 TC 208 TC 209 TC 344 K 1080 T 248 4.6 4.72 4.52 5.1 4.67
ŝij 0.10 0.01 -0.01 -0.02 0.03 r -0.22 -0.09 -0.25 0.19 -0.08
T 291 4.83 4.25 4.33 4.82 4.15 ŝij 0.06 -0.08 0.04 -0.27 r -0.22 0.01 0.23 0.08
TC 208 4.78 4.83 4.35 4.9 4.4 ŝij -0.04 0.09 -0.13 r -0.23 0.04 -0.03
TC 209 4.82 4.35 4.68 4.83 4.4 ŝij 0.00 0.06 r 0.14 -0.13
TC 344 4.92 4.8 5.08 4.85 4.98 ŝij 0.17 r -0.17
K 1080 4.63 4.03 4.32 4.57 4.92 ŝij r
Comparison of ŝij to ŝik - 0.034388 Comparison of ŝij to ŝkl - 0.022925
Given the high level of reciprocal effects of „nucleus-cytoplasm” interaction it is
necessary to test direct and reciprocal combinations in order to establish the best formulas
for obtaining high-oil hybrids
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CHAPTER VI
CONCLUSIONS
1. Analysis of maize germoplasm consisting of local populations, synthetic
populations and inbred lines of maize from ARDS Turda regarding protein and oil content
showed the existence of significant genetic variability that can be used to obtain corn
hybrids with high protein and oil content.
2. The variance of protein content in local populations of maize ranged between
11.20% and 15.60%, with an average of 13.69%, while oil content ranged between 3.80%
and 7.30%, with an average of 5.38%.
3. The variance of protein content in inbred maize lines ranged between 10.80% and
17.70%, with an average of 13.43%, as oil content ranged between 2.40% and 7.60%, with
an average of 4.16%.
4. Potential use in the increase of protein and oil content in maize were shown by the
following populations and inbred lines:
- local population Satu Lung/01(15.6% protein and 6.7% oil) and
Salva/01(15.5% protein and 7.1% oil)
- inbred lines :
- TC106(16.4% protein and 7,5% oil) and T442(15.6% protein and 7.2% oil)
inbred lines with flint kernels
- TC334(15.1% protein and 7.5% oil), T167(14.8% protein and 5.8% oil) and
T166(15.1% protein and 5.5% oil) inbred lines with flint × dent
- TC344A(15.2% protein and 7.6% oil), TB370(16% protein and 6.1% oil) şi
TC337(protein 15.5% and 5.9% oil) inbred lines with dent × flint kernels
- TC375(14.7% protein and 7.1% oil), TC353(15.1% protein and 5.6% oil) and
TD290(15.3% protein and 5.3% oil) inbred lines with dent kernels
5. In a balanced genetic system, with five inbred lines whose nucleus was transferred
by backcrossing (9-10) on four types of cytoplasms, in the heredity of protein and oil
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44
content genetic factors located in the nucleus, the cytoplasm and interaction between
factors located in cytoplasm and nucleus are involved.
6. In this balanced system, in the heredity of protein content, cytoplasm factors are
involved in a share of 7% and „nucleus-cytoplasm” interactions in a share of 57%. In
heredity of oil content cytoplasm factors have a minor contribution (1%), cytoplasm and
nucleus interactions having a contribution of 10%
7. In the balanced genetic system cytoplasm derived from the T248 line has led to an
increase of protein content in the isonuclear lines by 0.27%, in turn, cytoplasm derived
from TC221 inbred line reduced protein content by 0.29%. In both cases differences are
distinctly significant statistically.
8. Although the contribution of cytoplasm in heredity of oil content is reduced, the
cytoplasm of the TC177 inbred line has a significantly distinct positive contribution
(+0.31%) in improving oil content in the newly created isonuclear lines.
9. The nucleus transfer form the TB367 inbred line on the cytoplasm of the TC177 and
TB329 lines increased the oil content with 1.23% and 0.50%.
10. Study of protein and oil content of five sets of isonulcear inbred lines (each line
transferred on to six types of cytoplasm), under self pollinated or open pollinated
conditions revealed specific interactions between the type of cytoplasm and the transferred
nucleus to increase / decrease protein and oil content.
11. The results obtained for the protein content from open-pollinated kernels do not
always confirm the results obtained from self pollinated kernels. For this reason we believe
that self pollinated kernels give more accurate results regarding protein and oil content
when corn genotypes are cultivated on large surface areas, where opportunities for cross
pollination are reduced.
12. The oil content inheritance for the tested isonucleare inbred lines is influenced
primarily by the overall transmission capacity of the testers, by the genes with localization
in cytoplasm and „nucleus × cytoplasm” interaction. Share of variance for cataplasms
ranged between 1 and 58% for self pollinated ears and between 4 and 41% for open
pollinated ears.
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45
13. Heredity study of the protein content using a diallel system p (p-1) revealed the
importance of mean effects, general combining ability effect, specific combining ability
effects and the equally important effects of reciprocal interactions of „cytoplasm x
nucleus”. Maternal effects appear to be less important in the transmission of protein
content.
14. In the oil content heredity, studied in a diallel system on self pollinated and open
pollinated kernels revealed that additive gene effects, unaditive gene effects and reciprocal
interactions of „nucleus-cytoplasm” are important, maternal effects can be neglected.
15. Given the high level of reciprocal effects of „nucleus-cytoplasm” interaction it is
necessary to test direct and reciprocal combinations to determine the best hybrid
combinations to establish formulas for simple hybrids with high oil content.
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46
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