The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.)
The hybridological analysis was provided on several durum wheat genotypes with utilizing three F2 populations developed from the crossing between parental forms that differed in the invariable malt zone triplet on electrophoretic spectrum of α-amylase. Three components of this zone are controlled by...
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| Опубліковано в: : | Цитология и генетика |
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| Дата: | 2009 |
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Інститут клітинної біології та генетичної інженерії НАН України
2009
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| Цитувати: | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) / D.O. Prokopyk, M.Z. Antonyuk, T.K. Ternovskaya // Цитология и генетика. — 2009. — Т. 43, № 3. — С. 3-9 . — Бібліогр.: 21 назв. — англ. |
Репозитарії
Digital Library of Periodicals of National Academy of Sciences of Ukraine| _version_ | 1859665505424506880 |
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| author | Prokopyk, D.O. Antonyuk, M.Z. Ternovskaya, T.K. |
| author_facet | Prokopyk, D.O. Antonyuk, M.Z. Ternovskaya, T.K. |
| citation_txt | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) / D.O. Prokopyk, M.Z. Antonyuk, T.K. Ternovskaya // Цитология и генетика. — 2009. — Т. 43, № 3. — С. 3-9 . — Бібліогр.: 21 назв. — англ. |
| collection | DSpace DC |
| container_title | Цитология и генетика |
| description | The hybridological analysis was provided on several durum wheat genotypes with utilizing three F2 populations developed from the crossing between parental forms that differed in the invariable malt zone triplet on electrophoretic spectrum of α-amylase. Three components of this zone are controlled by three genes with an independent way of inheritance: one of them is located on the 6B or 5B chromosome, and two genes are located on the chromosomes of A subgenome.
Выполнен гибридологический анализ нескольких генотипов твердой пшеницы с использованием трех популяций F2, которые получены от скрещивания родительских форм, контрастных по электрофоретическим спектрам изозимов α амилазы в неизменном триплете мальт-зоны. Три компонента этой зоны контролируются тремя генами, наследующимися независимо: один ген хромосомы 6В и два гена хромосом субгенома А.
Виконано гібридологічний аналіз декількох генотипів твердої пшениці з використанням трьох популяцій F2, що одержані від схрещування батьківських форм, контрастних за електрофоретичними спектрами ізозимів α амілази в незмінному триплеті мальт-зони. Три комплекти цієї зони контролюються трьома генами, що успадковуються незалежно: один ген хромосоми 6В та два гени хромосом субгенома А.
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УДК 631.523:581.13
D.O. PROKOPYK, M.Z. ANTONYUK, T.K. TERNOVSKAYA
The National University «Kyiv�Mohyla Academy», Ukraine
THE GENETIC CONTROL
OF THE α�AMYLASE ISOZYMES
OF THE DURUM WHEAT
(TRITICUM DURUM DESF.)
The hybridological analysis was provided on several durum
wheat genotypes with utilizing three F2 populations developed
from the crossing between parental forms that differed in the
invariable malt�zone triplet on electrophoretic spectrum of α�
amylase. Three components of this zone are controlled by
three genes with an independent way of inheritance: one of
them is located on the 6B or 5B chromosome, and two genes
are located on the chromosomes of A subgenome.
Introduction. The cereal α�amylase genes are
divided into two large classes – AmyA and AmyB
[1]. The AmyA class, in its turn, is divided into two
subfamilies, Amy1 and Amy2 [2]. The AmyB class
contains the Amy3 subfamily [1, 3]. α�Amy�1 genes
are expressed in seed under the induction of gib�
berellic acid, which is produced by embryo [4], are
located on the chromosomes 6А, 6B, and 6D, and
encode a «malt» type of α�amylases that are
expressed at the initial stages of seed germination
(the 1–4th days). α�Amy�2 genes are located on 7А,
7B, and 7D chromosomes [5–8]. Expression of
these genes takes place on the later stages of germi�
nation (the 4–7th days), and their products are
regarded as a «green» type of amylases. α�Amy�
3 genes are expressed on the low level during the
embryo formation [2], but at the high one in the
aleurone layer during the germination [9, 10].
Amylase genes are multiple: we could distinguish
nearly 12–14 α�Amy�1 genes and 10–12 α�Amy�
2 ones [1, 11]. The difference between these
isozymes was revealed by means of isoelectric
focusing [7, 8]. α�Amy�3 genes are mapped on the
chromosomes of the 5th homeologous group [2,
12]. Both gene groups are polymorphic and usual�
ly vary among different varieties and landraces [8,
13, 14]. Nevertheless, this polymorphism is dis�
covered first of all by isoelectric focusing [7, 8, 14],
and native electrophoresis of amylase shows signif�
icantly less polymorphic spectra [5, 6, 15, 16].
Malt zone of common wheat (Triticum aestivum
L.) α�amylase electrophoretic spectrum in non�
denaturing PAAG usually consists of six compo�
nents. It was shown, that the first, second, and the
third components (in the direction of their mobili�
ty enhancement) are controlled by genes located
on 6D, 6B, and 6A chromosomes respectively [5,
6]. Genetic control of the next three bands of malt�
zone spectrum part is yet undetermined. The uti�
lization of nulli�tetrasomics for the 6th and the 7th
homeologous groups failed to dissect this triplet, as
all the bands were always present on the spectrum
[6, 16].
Previous investigations concerning genetic con�
trol of α�amylase spectrum components were con�
ducted on hexaploid and some tetraploid AbAbBB
wheat genotypes, and demonstrated a considerable
isozyme polymorphism [7, 8, 13, 14]. But in these
works the separation of isozymes was conducted by
means of isoelectric focusing in the thin�layer
PAAG, which is known for its labor� and resource�
intensivity, and, therefore, does not suit for genetic
ІSSN 0564–3783. Цитология и генетика. 2009. № 3 3
Оригинальные работы
© D.O. PROKOPYK, M.Z. ANTONYUK, T.K. TERNOVSKAYA,
2009
analysis. Moreover, this method, despite its high
resolution ability, did not shed light on the question
about genetic control of invariable malt�zone
bands. In previous works native electrophoresis did
not reveal any polymorphism in this zone among
common wheat genotypes [7, 8, 15, 16].
The question about localization of genes that
control permanent bands of the malt�zone is quite
important while these genes are suitable and efficient
chromosome�specific markers, which demonstrate
moderate polymorphism among common and
durum wheat genotypes. This paper expounds the
results of the investigation, the objective of which
was to ascertain the genetic control of durum wheat
α�amylase isozymes that form the invariable malt�
zone triplet.
Materials and methods. The genetic stocks used
in this research were forms and cultivars with the
genomic formula AABB, 2n = 28, namely durum
winter wheat cultivar Chornomor (developer –
O.I. Palamarchuk, Odessa), durum winter lines
Leucurum, Rubrum, and Candicance, kindly pro�
vided by Plant Groving Institute after Jurjiv V. Ja.,
(Kharkiv), winter durum wheat line Mutiko italikum
59h132, developed and kindly provided by breeder
V.V. Kostin, KNIAC, Krasnodar. The mentioned
parental forms were crossed after the diallelic
scheme without reciprocal crossings. The F1 plants
were allowed to self�pollinate under the isolators.
The F2 individuals were grown, and seeds obtained
from each plant were used in this investigation for
α�amylase extraction.
For α�amylase spectra obtainment the follow�
ing stocks were also used: common wheat cultivars
Chinese Spring, Kavkaz, Avrora (the genomic for�
mula AABBDD), and species T. boeoticum Boiss.
(AbAb), T. timоpheevii Zhuk. (AbAbGG), and nulli�
tetrasomics for 6th and 7th homologous groups,
developed from Chinese Spring cultivar, in all pos�
sible combinations.
The extraction buffer for α�amylase contained
sucrose 30 %, calcium chloride 0,2 %, brompheno�
lic blue stain. The PAAG for α�amylase was non�
denaturing, had a thickness 6,5 %, contained
0,02 M tris, 0,06 M glycine, and 5 M carbamide;
the ratio between acrilamide and bisacrilamide was
1 : 30. The electrode buffer has pH 8,4, and con�
tained 0,01 M tris and 0,06 M glycine. The α�amy�
lase pre�stain contained 12 g of hydrolysed starch,
134 ml 0,1 M acetic acid, and 3,6 g of sodium
acetate on the basis of 400 ml of solution. The
Lugol solution for the gel staining contains 0,03 M
potassium iodide, 0,01 M iodine, 8,4 % trichlo�
roacetic acid. The α�amylase isozymes were
extracted from the germinated seeds with coleop�
tiles 0,5–3 cm long. The germinated seeds were
crashed in 1 ml of extraction buffer and incubated
during 6–18 h at 4 °С. Before drifting on the gel
the samples were centrifuged for 3 min under the
acceleration 8000 xg; on each cell 5 μl of extract
were drifted [16]. The α�amylase separation was
performed from cathode to anode under the volt�
age 300 V and the current strength 0,4–1 mA for
each centimetre of gel. After the run gels were incu�
bated in the 3 % starch solution during 40–60 min,
washed a little and stained with Lugol solution.
Results and discussion. On the elecrtophoretic
spectra of α�amylase derived from common and
durum wheat genotypes (Fig. 1) components
1–6 correspond to malt zone, and 7–9 – to the
green one [7, 8, 16]. Band 1 is controlled by Amy�
D1 gene, located on 6D chromosome [7, 16]. This
explains why the spectrum of durum wheat lacks it.
Bands 2 and 3 are the products of α�Amy�1 genes,
which are located on the chromosomes 6B and 6A
respectively [7, 16]. Components 4, 5, and 6 form
so named invariable malt�zone triplet, and the
genetic control of the respective isozymes is yet
undetermined. Bands 7, 8, and 9 of common
wheat and 8, 9 of durum wheat are the products of
α�Amy�2 genes, which are located on the chromo�
somes of the 7th homeologous group 7D, 7A, and
7B and 7A, 7B respectively [7, 16].
The polymorphism in the invariable triplet was
observed among five durum wheat genotypes (Fig.
1, a–е). The spectrum of Leucurum lacks band 4
(Fig. 1, c), and demonstrates single bands 5 and 6,
whereas the spectrum of MI (Fig. 1, d) has band 4,
and shows double bands 5 and 6. They are numer�
ated as 5а, 5b, and 6а, 6b. The invariable triplet
spectra of Rubrum and Candicance have all three
components (Fig. 1 a, e), and Chornomor’s spec�
trum (Fig. 1, b) has bands 4, 6, and double 5. The
availability of the polymorphism in the malt�zone
triplet among some durum wheat representatives
provides a possibility of using these genetic stocks
in the genetic analysis of the triplet’s components
genetic control.
The electrophoretic spectra of different nulli�
tetrasomics for the 6th homeologuos group approve
ISSN 0564–3783. Цитология и генетика. 2009. № 34
D.O. Prokopyk, M.Z. Antonyk, T.K. Ternovskaya
previous results and provide evidence that whichev�
er chromosome was absent, none of the triplet’s
bands disappeared. So, we may conclude, first of all,
that isozymes, which genes locate on different
chromosomes, may have the same electrophoretic
mobility and therefore collide on the spectrum. To
check this assumption, we examined the spectra of
the diploid wheat T. boeoticum Boiss. (АbАbgenome)
and tetraploid wheat T. timopheevi Zhuk.
(AbAbGG genome).
The spectrum of diploid species T. boeoticum is
presented only by two bands, which are both located
in the invariable triplet zone. These bands are the 5th
and the 6th, and the latter could vary among different
einkorn samples and be either single or double (Fig.
2, a, b). Such a spectrum could appear in two cases:
when Amy�A1 gene is a cluster and its different alle�
les could provide two or three electrophoretic bands,
or when the genome of einkorn has two distinct
genes, which products form the invariable malt�zone
triplet. The latter case also provides two alternatives,
when both genes are located on the 6A, or when one
of them is Amy3, which is located on the 5A chromo�
some. This might explain why the exploitation of
nulli�tetrasomics for the 6th group is ineffective in the
localization of genes that control malt�zone bands.
The spectrum of the einkorn lacks bands 3 and 8,
which are encoded by Amy�A1 and Amy�A2 genes
located on chromosomes 6А and 7А respectively of
common (AuAuBBDD) and, possibly, durum wheat.
So, the α�amylase spectrum of einkorn wheat gives
an insight into which bands are produced exception�
ally by Ab genome, which is present as a subgenome
in the tetraploid T. timopheevii. It differs from the
spectrum provided by Au subgenome, which we could
observe on spectra of common and durum wheats.
The α�amylase spectrum of T. timopheevii lacks
bands 3 and 8, which are present on spectra of com�
mon and durum wheat and point on the expression
of genes located on chromosomes 6A and 7A. So,
there is some accordance between the spectra of
einkorn and Timopheevii wheat. The comparison
of diploid and tetraploid wheat spectra leads to a
conclusion that the upper component (4) on the
invariable triplet and the lower band of the green�
zone are controlled by chromosomes of G and B
subgenomes of Timopheevii and durum wheats,
respectively.
Wild einkorn, the boeoticum wheat, also lacks on
its spectrum bands 3 and 7, which are controlled by
chromosomes of common wheat Au subgenome.
Therefore, we could presume that the initial gene
of the einkorn wheat of the Ab subgenome is dupli�
cated in comparison to genome Аb, and these
genes encode different bands. So, the comparison
ІSSN 0564–3783. Цитология и генетика. 2009. № 3 5
The genetic control of the α�amylase isozymes of the durum wheat (Triticum durum Desf.)
Fig. 1. The electrophoretic spectra of some durum and com�
mon wheat genotypes: a – Rubrum, b – Chornomor, c – Leu�
curum, d – Mutiko italicum, e – Candicance, f – Avrora,
g – Kavkaz
Fig. 2. The electrophoretic spectra of α�amylase isozymes
of some T. timopheevii and T. boeoticum genotypes: a, b –
T. boeoticum, c – T. timopheevii, d – Avrora; e – Mutiko
italicum
of spectra of diploid, two tetraploid and one hexa�
ploid wheat provides evidence that the upper band
of the triplet is encoded by G (B) subgenome, but
leaves unclear how the next two bands are con�
trolled. The other assumption occurs: if two bands
of triplet, 5 and 6, are controlled by one genome
(Аb of T. boeoticum), it might be possible that B or
G subgenomes, as well as D one of common wheat,
may encode two isozymes each. If some isozymes
have equal electrophoretic mobility, they should
collide on the spectrum and as a result decrease the
number of the bands on spectrum and form the
invariable triplet. Moreover, that should cause inef�
fectiveness of nulli�tetrasomics exploitation in gene
location definition. Fig. 3 demonstrates two most
plain schemes of electrophoretic band genetic con�
trol in hexaploid wheat. It should be noted that nei�
ther of these schemes for common wheat, a and b,
is more trustworthy than the other due to the
absence of the appropriate genetic stocks.
It must not be ruled out that one of the α−amy�
lase bands of the T. boeoticum is controlled by Amy�
A1 (6А), and the other is controlled by Amy�A3
(5А). Fig. 4 demonstrates six most plain schemes
of electrophoretic band genetic control in tetra�
ploid wheat, which explain the presence of three
components in the invariable spectrum part. Durum
wheat genotypes, which were used in our investiga�
tion and showed the polymorphism in the invari�
able malt�zone triplet, offer an opportunity of shed�
ding some light on the genetic control of the invari�
able triplet bands.
For the genetic analysis the combinations Leu�
curum � Rubrum, MI � Leucurum, and MI � Ru�
brum were used due to their difference in the α�
amylase malt�zone triplet on the electrophoretic
spectra (Fig. 1). Four seeds from each F2 individ�
ual were randomly taken, and the extracted α�
amylase was separated electrophoretically. On the
basis of the four seeds spectra the genotype for
ISSN 0564–3783. Цитология и генетика. 2009. № 36
D.O. Prokopyk, M.Z. Antonyk, T.K. Ternovskaya
Fig. 3. Some variants of chromosomal control of the bands in invariable triplet of α�amylase in common wheat
Fig. 4. Some variants of chromosomal control of the bands in invariable triplet of α�amylase in durum wheat
The distribution of F2 individuals into phenotypic classes after α�amylase
electrophoretic spectrum bands in the analyzed combinations
Phenotypic
classes
of empiric classes of theoretic classes
Phenotypic
classes
of empiric classes of theoretic classes
Leucurum � MI Leucurum � Rubrum
MI � Rubrum
27 M1M2M3
9 M1M2L3
9 M1L2M3
9 L1M2M3
3 M1L2L3
3 L1L2M3
3 L1M2L3
1 L1L2L3
Total
χ2 value
65
19
18
17
8
2
12
3
144
60,75
20,25
20,25
20,25
6,75
6,75
6,75
2,25
9,05 < χ2
st 0,01
3 R1
1 L1
Total
χ2 value
9 M2M3
3 M2R3
3 R2M3
1 R2R3
48
21
31
12
112
63
21
21
7
20,96 > χ2
st 0,01
76
23
99
74.25
24.25
0,16< χ2
st 0,01
Volumes Volumes
genes that control α�amylase of each F2 individual
was reconstructed. As cultivar Leucurum has no
upper band on its spectrum, we can not distinguish
a homozygote for its alternative allele, which
demonstrates a band, from a heterozygote for this
gene. So, there could be only two phenotypic class�
es: homozygotes for the null�allele (no band) and
general class that consists of homo� and heterozy�
gotes, demonstrating the upper band.
The same concerns the second and the third
bands of triplet. The double bands provided by MI
genotype collide with the respective single bands of
other genotypes; therefore, according to genes that
control two lower triplet bands F2 plants could be
divided into two phenotypic classes: homozygotes
with the single band phenotype, and homo� and
heterozygotes with the double bands phenotype.
From the α�amylase electrophoretic spectra it
could be drawn that the individuals from the cross
combination Leucurum (L) � MI (M) differ in three
genes. We expect 3 : 1 segregation for one gene (the
band is present or it is absent); consequently, for
three genes phenotypic ratios should match (3 : 1)3.
In the cross combination Leucu�rum � Rubrum
(R), where parental forms differ in one gene, which
encodes the upper triplet band, 3 : 1 segregation is
expected. MI and Rubrum differ in two genes
controlling α�amylase isozymes, therefore, in F2
(3 : 1)2 ratio between phenotypic classes is expect�
ed. Fig. 5 demonstrates electrophoretic spectra of
F3 individuals obtained from the crossing different
cultivars; bands are defined via letter (that means
corresponding cultivar) and number (that means
the position in the triplet zone). Bands M1 and
ІSSN 0564–3783. Цитология и генетика. 2009. № 3 7
The genetic control of the α�amylase isozymes of the durum wheat (Triticum durum Desf.)
Fig. 5. Some examples of α�amylase electrophoretic spectra of four F3 seeds grown on the F2
plant, crossing combination MI � Leucurum: a – M1L1M2L2M3L3 genotype of the parent
plant F2; b – M1L1L2L2M3M3; c – M1L1L2L2L3L3; d – L1L1L2L2M3M3
Fig. 6. Some examples of α�amylase electrophoretic spectra of four F3 seeds grown on the F2
plant, crossing combination Leucurum � Rubrum: a – L1L1 genotype of the parent plant F2;
b–d – L1R1
R1, R2 and L2, R3 and L3 are not analyzed due to
their identity. Bands M1 and R1 dominate over L1;
consequently, M1M1 and M1L1, R1R1 and
R1L1 are identical, too. In the same way, band
М2 dominates over R2, and М3 – R3, and geno�
types М2М2 and М2R2, М3М3 and М3R3 could
not be distinguished.
The distribution of the F2 plants into phenotyp�
ic classes (Table) agrees with the theoretically
expected one in those two crossing combinations,
where Leucurum was one of the parental forms.
The empiric distribution differs from the theoretic
one in the combination, in which MI was one of
the parental forms. A deficit of plants in classes
with bands M2 and M3 is observed. MI has rather
weak frost�resistance, so, it is fear to presume that
among F2 individuals did not winter those ones
that possessed, first of all, originated from MI 5A
chromosome, which is known as a carrier of sever�
al genes that control frost resistance [17–19].
Under negative selection occurred genes located
on the 5A chromosome, among which was Amy�
A3 gene. Consequently, we could presume that the
central band of the invariable malt�zone triplet is
controlled exactly by this gene. So, schemes d and
f in Fig. 4 could be thought of ones as more realis�
tic as for chromosomal control of the bands in the
invariable part of α�amylase spectrum of durum
wheat.
Conclusion. The invariable malt�zone triplet of
α�amylase in tetraploid wheats with genome for�
mulas AuAuBB and AbAbGG is controlled by three
genes, which recombine independently. Two of them
are controlled by the genes of the A subgenome:
the lowest band is a product of the Amy�A1, locat�
ed on the 6A chromosome, the central triplet’s
band is a product of Amy�A3, located on 5A chro�
mosome. The upper band is encoded by subgenomes
B of T. durum and G of T. timopheevaii.
Д.А. Прокопик, М.З. Антонюк, Т.К. Терновская
ГЕНЕТИЧЕСКИЙ КОНТРОЛЬ ИЗОФЕРМЕНТОВ
α�АМИЛАЗЫ У ТВЕРДОЙ ПШЕНИЦЫ
(TRITICUM DURUM DESF.)
Выполнен гибридологический анализ нескольких
генотипов твердой пшеницы с использованием трех
популяций F2, которые получены от скрещивания ро�
дительских форм, контрастных по электрофоретичес�
ким спектрам изозимов α�амилазы в неизменном
триплете мальт�зоны. Три компонента этой зоны кон�
тролируются тремя генами, наследующимися незави�
симо: один ген хромосомы 6В и два гена хромосом
субгенома А.
Д.А. Прокопик, М.З. Антонюк, Т.К. Терновська
ГЕНЕТИЧНИЙ КОНТРОЛЬ ІЗОФЕРМЕНТІВ
α�АМІЛАЗИ У ТВЕРДОЇ ПШЕНИЦІ
(TRITICUM DURUM DESF.)
Виконано гібридологічний аналіз декількох геноти�
пів твердої пшениці з використанням трьох популяцій
F2, що одержані від схрещування батьківських форм,
контрастних за електрофоретичними спектрами ізо�
зимів α�амілази в незмінному триплеті мальт�зони.
Три комплекти цієї зони контролюються трьома гена�
ми, що успадковуються незалежно: один ген хромосо�
ми 6В та два гени хромосом субгенома А.
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Received 17.07.08
ІSSN 0564–3783. Цитология и генетика. 2009. № 3 9
The genetic control of the α�amylase isozymes of the durum wheat (Triticum durum Desf.)
|
| id | nasplib_isofts_kiev_ua-123456789-66637 |
| institution | Digital Library of Periodicals of National Academy of Sciences of Ukraine |
| issn | 0564-3783 |
| language | English |
| last_indexed | 2025-11-30T10:56:42Z |
| publishDate | 2009 |
| publisher | Інститут клітинної біології та генетичної інженерії НАН України |
| record_format | dspace |
| spelling | Prokopyk, D.O. Antonyuk, M.Z. Ternovskaya, T.K. 2014-07-19T18:12:38Z 2014-07-19T18:12:38Z 2009 The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) / D.O. Prokopyk, M.Z. Antonyuk, T.K. Ternovskaya // Цитология и генетика. — 2009. — Т. 43, № 3. — С. 3-9 . — Бібліогр.: 21 назв. — англ. 0564-3783 https://nasplib.isofts.kiev.ua/handle/123456789/66637 631.523:581.13 The hybridological analysis was provided on several durum wheat genotypes with utilizing three F2 populations developed from the crossing between parental forms that differed in the invariable malt zone triplet on electrophoretic spectrum of α-amylase. Three components of this zone are controlled by three genes with an independent way of inheritance: one of them is located on the 6B or 5B chromosome, and two genes are located on the chromosomes of A subgenome. Выполнен гибридологический анализ нескольких генотипов твердой пшеницы с использованием трех популяций F2, которые получены от скрещивания родительских форм, контрастных по электрофоретическим спектрам изозимов α амилазы в неизменном триплете мальт-зоны. Три компонента этой зоны контролируются тремя генами, наследующимися независимо: один ген хромосомы 6В и два гена хромосом субгенома А. Виконано гібридологічний аналіз декількох генотипів твердої пшениці з використанням трьох популяцій F2, що одержані від схрещування батьківських форм, контрастних за електрофоретичними спектрами ізозимів α амілази в незмінному триплеті мальт-зони. Три комплекти цієї зони контролюються трьома генами, що успадковуються незалежно: один ген хромосоми 6В та два гени хромосом субгенома А. en Інститут клітинної біології та генетичної інженерії НАН України Цитология и генетика Оригинальные работы The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) Генетический контроль изоферментов α-амилазы у твердой пшеницы (triticum durum Desf.) Генетичний контроль ізоферментів α-амілази у твердої пшениці (triticum durum Desf.) Article published earlier |
| spellingShingle | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) Prokopyk, D.O. Antonyuk, M.Z. Ternovskaya, T.K. Оригинальные работы |
| title | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) |
| title_alt | Генетический контроль изоферментов α-амилазы у твердой пшеницы (triticum durum Desf.) Генетичний контроль ізоферментів α-амілази у твердої пшениці (triticum durum Desf.) |
| title_full | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) |
| title_fullStr | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) |
| title_full_unstemmed | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) |
| title_short | The genetic control of the α-amylase isozymes of the durum wheat (triticum durum Desf.) |
| title_sort | genetic control of the α-amylase isozymes of the durum wheat (triticum durum desf.) |
| topic | Оригинальные работы |
| topic_facet | Оригинальные работы |
| url | https://nasplib.isofts.kiev.ua/handle/123456789/66637 |
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