**3.2 Gene encoding starch biosynthesis (waxy genes)**

Polymorphisms of *Wx* gene markers that were found of 8 polymorphisms alleles and 14 genotypes of the *Wx* genes [16]. The polymorphism detected in sago palm was in line of the polymorphism in *Triticum aestivum* L. by using the *Wx* (SunI) gene markers [14]. The number of alleles and genotypes of sago palm at the level populations and islands varies as well as their frequency [17]. The Wx gene variations found in sago are similar with the *Wx* gene variations on wheat [15]. Furthermore, Scientist were reported a high Wx gene polymorphism in barley [9] and in rice [13]. This phenomenon indicates that the source of the *Wx* genes diversity is the Papua islands Papua and Sulawesi islands because these islands are found genotypes that are not found on other islands [16]. If the center of diversity is the object of attention, then the island of Papua is the center of diversity of the *Wx* genes because the most genotypes of *Wx* genes are found on the island of Papua [17].

The genetic diversities of *Wx* genes that was observed to the sago palm from various islands were shown varied. The genetic diversities calculation results showed varying values except for samples from Jawa [17]. The sago palm variations were detected, those a reflection of sago palm variations that it occurs in the several islands in Indonesia [16]. The *Wx* gene is one of many genes that it is regulated biosynthesis process for resulting starch of plants, including sago palm. If the *Wx* gene has high variations that will be resulting various quantity and quality of starch. In the previous studies were reported the quantities of starch accumulation of sago palm range from 28 to 710 kg trunk−1 [45] and starch accumulation of sago palm trunk−1 will be depend on the varieties [21]. The *Wx* gene was one of the genes that influenced of starch synthesis in rice endosperm [54]. Two alleles of the Wx gene that is Wxa and Wxb gene were reported regulating to increase *Wx* protein and amylose content [10]. *Wx* allelic pulp in wheat showed a significantly different reduction in amylose content [12] and recombinant inbred line (RIL) of wheat that has integrated three *Wx* genes in their genome was reported resulting high quality starch than wheat RIL which did not contain the three *Wx* genes [32].

The heterozygote values of sago palm in the populations that was observed were shown variation from 0.52–1.00 with a low standard deviation of 0.0000– 0.0014 [17]. The heterozygosity variations were indicated variations in the *Wx* gene in the genome of sago palm. The key gene that influences starch synthesis in rice endosperm is the *Wx* gene [54]. Variation of the *Wx* gene causes a variation in the viscosity of starch production in wheat (Boggini et al. 2001). The *Wx*a and

**35**

*Adaptive Evolution and Addressing the Relevance for Genetic Improvement of Sago Palm…*

*Wx*b alleles were found to regulate quantitative levels of *Wx* protein as well as

The genetic hierarchy calculation using AMOVA shows that individuals and populations was estimated significantly different [17]. The differentiation values based on the chi-square test at the population and island level were found sago palm differentiated that occurs at the population and island level [17]. The detected variance is an indicator that the *Wx* gene varies both at the individual level and at the population level. Previous studies of sago palm using different markers also showed that sago palm varied both in terms of quantity and quality of production [21, 45]. The allelic levels of *Wx* genes and their interactions in starch-producing plants were reported increasing quality and quantity of starch production [10, 11, 32], and [55]. It is predicted that the *Wx* gene variation in sago palm is one of the genes that determines the variation in the quantity and quality of sago starch yields [16]. The sample diversity at the inter-island population was not significantly different based on the AMOVA value as was the sample at the inter-island population. This phenomenon indicates that the variation of the *Wx* gene in sago palm is more caused by variations at the individual and populations, not due to the isolation of different distances and geographic differences due to

Genetic relationship among individual shows that sago palm are grouped into four groups based on dendrogram construction [17]. The division into four groups was strengthened by the MDS test which showed the sample was distributed in four quadrants. The data illustrates that certain individuals are not grouped based on population origin but rather mixed with each other with different population origins and different local names [16]. This description implies that local names are not appropriate when used as a reference for determining the number of species or varieties of sago palm without the support of other data such as molecular data. In the vicinity of Sentani Lake, the local community revealed that there were 21 types of sago palm based on morphology and scientist found only 15 species based on the

Genetic relationship of sago palm in the population level shows that sago palm from the populations of Jayapura, Serui, Sorong, and Pontianak are closely related and form group I, samples from populations from Manokwari, Palopo, and Selat Panjang cluster to form group II, then groups III and IV only formed from one population. The grouping of the population into four groups is also strengthened by the MDS test which shows the population sample is distributed in four quadrants [16]. Previous studies have discussed the genetic relationships of populations using various markers [39, 47, 49, 51]. Populations contained in one group are closely related, on the other hand, populations in different groups are not closely related. The differences in a population is thought to be caused by outbreeding so that the population experiences differentiation. Population differentiation can be caused by pollen migration [56]. In general, it can be interpreted that there is a tendency for sago palm in Indonesia to be differentiated inter-island and among island based on the *Wx* gene marker. Differentiation can be caused by evolutionary processes, georaphic isolation, distance isolation, genetic drift and gene flow. Population differentiation is caused by evolution, natural selection, migration, and genetic drift [57] and the differentiation of

Based on cpDNA markers, various polymorphic and haplotypic alleles were found in sago palm. Studies related to the use of the NTCP21 and NTCP22 markers

*DOI: http://dx.doi.org/10.5772/intechopen.94395*

the low FCT value of 0.06044 [17].

Cruciferae due to gene flow [58].

**3.3 Chloroplast DNA (cpDNA) marker**

same marker [21].

amylose content [10].

*Adaptive Evolution and Addressing the Relevance for Genetic Improvement of Sago Palm… DOI: http://dx.doi.org/10.5772/intechopen.94395*

*Wx*b alleles were found to regulate quantitative levels of *Wx* protein as well as amylose content [10].

The genetic hierarchy calculation using AMOVA shows that individuals and populations was estimated significantly different [17]. The differentiation values based on the chi-square test at the population and island level were found sago palm differentiated that occurs at the population and island level [17]. The detected variance is an indicator that the *Wx* gene varies both at the individual level and at the population level. Previous studies of sago palm using different markers also showed that sago palm varied both in terms of quantity and quality of production [21, 45]. The allelic levels of *Wx* genes and their interactions in starch-producing plants were reported increasing quality and quantity of starch production [10, 11, 32], and [55]. It is predicted that the *Wx* gene variation in sago palm is one of the genes that determines the variation in the quantity and quality of sago starch yields [16]. The sample diversity at the inter-island population was not significantly different based on the AMOVA value as was the sample at the inter-island population. This phenomenon indicates that the variation of the *Wx* gene in sago palm is more caused by variations at the individual and populations, not due to the isolation of different distances and geographic differences due to the low FCT value of 0.06044 [17].

Genetic relationship among individual shows that sago palm are grouped into four groups based on dendrogram construction [17]. The division into four groups was strengthened by the MDS test which showed the sample was distributed in four quadrants. The data illustrates that certain individuals are not grouped based on population origin but rather mixed with each other with different population origins and different local names [16]. This description implies that local names are not appropriate when used as a reference for determining the number of species or varieties of sago palm without the support of other data such as molecular data. In the vicinity of Sentani Lake, the local community revealed that there were 21 types of sago palm based on morphology and scientist found only 15 species based on the same marker [21].

Genetic relationship of sago palm in the population level shows that sago palm from the populations of Jayapura, Serui, Sorong, and Pontianak are closely related and form group I, samples from populations from Manokwari, Palopo, and Selat Panjang cluster to form group II, then groups III and IV only formed from one population. The grouping of the population into four groups is also strengthened by the MDS test which shows the population sample is distributed in four quadrants [16]. Previous studies have discussed the genetic relationships of populations using various markers [39, 47, 49, 51]. Populations contained in one group are closely related, on the other hand, populations in different groups are not closely related. The differences in a population is thought to be caused by outbreeding so that the population experiences differentiation. Population differentiation can be caused by pollen migration [56]. In general, it can be interpreted that there is a tendency for sago palm in Indonesia to be differentiated inter-island and among island based on the *Wx* gene marker. Differentiation can be caused by evolutionary processes, georaphic isolation, distance isolation, genetic drift and gene flow. Population differentiation is caused by evolution, natural selection, migration, and genetic drift [57] and the differentiation of Cruciferae due to gene flow [58].

#### **3.3 Chloroplast DNA (cpDNA) marker**

Based on cpDNA markers, various polymorphic and haplotypic alleles were found in sago palm. Studies related to the use of the NTCP21 and NTCP22 markers

*Genetic Variation*

the island of Java are separate from other islands based on the RAPD marker [38]. Here there is something interesting to observe because the sample at the island level forms a group together with samples from other islands that are far away, such as the sample from the island of Papua which forms a group together with the sample from the island of Sumatra. When examined from the migration side, it is possible that individual sago palm from Papua population have mingled with sago palm from the island of Sumatra. This phenomenon is possible because the molecular markers (RAPD) used are not as conservative as the cpDNA molecular markers that are uni-parental inherited [24, 25]. The RAPD marker is a nucleus genomic molecular marker associated with the DNA recombination process and is biparental inherited [39] so that the RAPD marker is a molecular marker that has a relatively short conservative time (one generation) compared to the cpDNA molecular marker. Previous studies suggest that higher variation is found using nucleus genomic

Polymorphisms of *Wx* gene markers that were found of 8 polymorphisms alleles and 14 genotypes of the *Wx* genes [16]. The polymorphism detected in sago palm was in line of the polymorphism in *Triticum aestivum* L. by using the *Wx* (SunI) gene markers [14]. The number of alleles and genotypes of sago palm at the level populations and islands varies as well as their frequency [17]. The Wx gene variations found in sago are similar with the *Wx* gene variations on wheat [15]. Furthermore, Scientist were reported a high Wx gene polymorphism in barley [9] and in rice [13]. This phenomenon indicates that the source of the *Wx* genes diversity is the Papua islands Papua and Sulawesi islands because these islands are found genotypes that are not found on other islands [16]. If the center of diversity is the object of attention, then the island of Papua is the center of diversity of the *Wx* genes because the most genotypes of *Wx* genes are found on the island of

The genetic diversities of *Wx* genes that was observed to the sago palm from various islands were shown varied. The genetic diversities calculation results showed varying values except for samples from Jawa [17]. The sago palm variations were detected, those a reflection of sago palm variations that it occurs in the several islands in Indonesia [16]. The *Wx* gene is one of many genes that it is regulated biosynthesis process for resulting starch of plants, including sago palm. If the *Wx* gene has high variations that will be resulting various quantity and quality of starch. In the previous studies were reported the quantities of starch accumulation of sago palm range from 28 to 710 kg trunk−1 [45] and starch accumulation of sago palm trunk−1 will be depend on the varieties [21]. The *Wx* gene was one of the genes that influenced of starch synthesis in rice endosperm [54]. Two alleles of the Wx gene that is Wxa and Wxb gene were reported regulating to increase *Wx* protein and amylose content [10]. *Wx* allelic pulp in wheat showed a significantly different reduction in amylose content [12] and recombinant inbred line (RIL) of wheat that has integrated three *Wx* genes in their genome was reported resulting high quality starch than wheat RIL which did not

The heterozygote values of sago palm in the populations that was observed were shown variation from 0.52–1.00 with a low standard deviation of 0.0000– 0.0014 [17]. The heterozygosity variations were indicated variations in the *Wx* gene in the genome of sago palm. The key gene that influences starch synthesis in rice endosperm is the *Wx* gene [54]. Variation of the *Wx* gene causes a variation in the viscosity of starch production in wheat (Boggini et al. 2001). The *Wx*a and

markers rather than cpDNA markers [39–41, 53].

**3.2 Gene encoding starch biosynthesis (waxy genes)**

**34**

contain the three *Wx* genes [32].

Papua [17].

in potato have also demonstrated allele polymorphisms in potato [59]. Locus rpl1671, NTCP21, and NTCP22 on sago were detected in three haplotypes out of 10 haplotypes which were specific haplotypes in populations from Jayapura and one specific haplotype each for populations from Serui, Palopo, and Pontianak [26]. The specific haplotype phenomenon is also found in several types of plants i.e. Cunninghamia spp. [60], *Pinus sylvestris* L. [27], and *Alyssum* spp. [29]. The specific haplotypes were found in a population, those indicated the source of diversities in a population. The specific haplotypes of sago palm were found in the populations of Papua, Sulawesi, and Kalimantan indicated the provenance of the diversities, while the most haplotypes of sago palm diverse is the population from Jayapura then followed by the sago palm population from Serui [17]. The large number of haplotypes reflects the high variation in a population in line of the *Abies alba* Mill population [28]. The differences in chloroplast haplotypes in each population reflect differences in genetic entities or sources of variation [29]. The number of haplotypes that were found to be present together in each population is an indication that genetic similarities among individual in a population. It is hypothetically that the sago palm migration by carrying of people. Four haplotypes of 10 haplotypes of sago palm were found in to two or more populations, which means that only four haplotypes were found migration through various kinds of intermediaries. The same thing was also found in *P. sylvestris* L. and *A. alba* Mill. referred to as the refugial population [27, 28].

Population genetic diversity shows that the population from Papua has the highest number of haplotypes (∑H), the number of polymorphic sites (S), and the highest percentage of haplotype polymorphics compared to other populations. A value (HE) equal to one means that no haplotype numbers are the same in individual samples in a population (single haplotype individuals) as happened in the population from Bogor. This is similar with individual haplotype on *P. sylvestris* L [27]. Previous studies on sago palm using RAPD markers showed that sago plant diversity at the individual level was also high [48, 52].

The genetic hierarchy based on cpDNA was estimated by using analysis of molecular variance (AMOVA) was calculated of differentiation level of population samples at the inter-island level (−3.88% and FCT = −0.03884), between populations within islands (8.49% and FSC = 0.08177), and Papua and others (5.05% and FCT 0.05054) which is low with the probability value not significantly different. High percentage values of variance were observed at the level among individuals (95.39% and FST = 0.04610) and between populations (5.91% and FST = 0.05914) with significantly different probability values [26]. The same thing was also found in *P. sylvestris* L., namely the percentage value of variance between populations (3.24%) with a significantly different probability [27]. The negative value observed at the inter-island level indicates that the sample island level does not contribute to the total measured variance. This phenomenon resembles the tetraploid alfalfa population [47]. Negative correlation coefficients have a biological significance in that the samples at the inter-island level are more closely related than those at the island level [61]. Based on this, it indicates that island or geographic differences do not cause variations in the chloroplast genome, even though the distance between one island and another is far (hundreds to thousands of kilometers). The variation between individuals and between populations contributed 95.39% and 5.91% to the total variety and was significantly different [18]. The results observed were similar with Abies species that was only a small variance value between populations (6.10%), high proportion of variance within the population or between individuals (74.66%) [62]. A low trend of genetic variability between populations is also found in Pinaceae [39] and species other than pine [63].

**37**

on apple plants [24].

*Adaptive Evolution and Addressing the Relevance for Genetic Improvement of Sago Palm…*

Genetic differentiation based on the Fst value shows that among the populations being compared, only the population from Jayapura is significantly different from the population from Palopo and Pontianak [26]. These populations based on cpDNA markers each have a genetic entity, which means that the diversity that occurs in this population has appeared separately since ancient times (thousands of years ago). The genetic differentiation of samples at the

Jayapura is different from the population originating from Serui, Manokwari, Sorong, Palopo, Pontianak, and Selat Panjang and the population originating from Serui is different from the population originating from Pontianak but not

level, it indicates that the source of sago plant diversity at the population level is the population from Jayapura, Serui, and Pontianak. Genetic differentiation

ing from the island of Papua are different from samples from the islands of Sulawesi and Kalimantan [26]. This is in line with the specific haplotypes found on the three islands. For this reason, it is suggested that the sources of sago palm diversity based on the samples tested are the islands of Papua, Sulawesi and Kalimantan. This data is also consistent with the grouping of sago palm samples through phylogenetic construction at the island level which divides the samples into three groups, namely the Papua group, the Sulawesi group and the Kalimantan group. This indicates that the source of the diversity of sago palm in Indonesia, apart from being on the island of Papua, is also found in other islands,

The genetic relationship of the samples at the individual level shows that the samples are classified into three groups based on the phylogenetic construction. Sago palm relationship studies previously show that sago palm originating from the Malay Archipelago and several samples of sago from Indonesia clustered into two groups and two sub-groups based on the RAPD markers [48]. The sago relationship study focused on the Indonesian archipelago, but with a larger number of samples, showed that the sample individuals were divided into four groups based on RAPD markers [52]. The discrepancy in the division of the number of groups (groups) between the groupings based on cpDNA markers and RAPD markers that was observed in previous studies is something that is often found in studies of genetic relationship using molecular markers. Previous genetic related studies which showed that different molecular markers led to different groupings of certain plants by using cpDNA, RAPD and isozyme markers in Pseudotsuga spp. (Pinaceae) [39], cpDNA and inter-SSR (ISSR) markers in the nucleus genome on *Brassica oleraceae* L. plants using [41], and using cpDNA and mitochondrial DNA

Based on cpDNA, the sources of sago diversity in Indonesia are predicted to come from three islands, namely Papua, Sulawesi and Kalimantan. It is suspected that from these three islands, individual sago palm experienced migration in line with migration and population mobilization in Indonesia that had occurred hundreds of years ago. This assumption is reinforced by haplotype data, phylogenetic analysis, and genetic hierarchies which show that samples at the inter-population level and between individuals are significantly different, which means that there are one or more different individuals or populations. Although the source of diversity is found in three islands, if the number of haplotypes is the size of the center of diversity, then the island of Papua is the center of diversity of sago palm in Indonesia because that island is found in the largest number of haplotypes compared to other islands. Apart from the highest number of haplotypes, on the island of Papua, the wild relatives of sago palm

test shows that the population originating from

test at the population

test shows that samples originat-

*DOI: http://dx.doi.org/10.5772/intechopen.94395*

population level based on the X<sup>2</sup>

different from other populations [17]. Based on the X<sup>2</sup>

of samples at the island level based on the X<sup>2</sup>

namely Sulawesi and Kalimantan [26].

#### *Adaptive Evolution and Addressing the Relevance for Genetic Improvement of Sago Palm… DOI: http://dx.doi.org/10.5772/intechopen.94395*

Genetic differentiation based on the Fst value shows that among the populations being compared, only the population from Jayapura is significantly different from the population from Palopo and Pontianak [26]. These populations based on cpDNA markers each have a genetic entity, which means that the diversity that occurs in this population has appeared separately since ancient times (thousands of years ago). The genetic differentiation of samples at the population level based on the X<sup>2</sup> test shows that the population originating from Jayapura is different from the population originating from Serui, Manokwari, Sorong, Palopo, Pontianak, and Selat Panjang and the population originating from Serui is different from the population originating from Pontianak but not different from other populations [17]. Based on the X<sup>2</sup> test at the population level, it indicates that the source of sago plant diversity at the population level is the population from Jayapura, Serui, and Pontianak. Genetic differentiation of samples at the island level based on the X<sup>2</sup> test shows that samples originating from the island of Papua are different from samples from the islands of Sulawesi and Kalimantan [26]. This is in line with the specific haplotypes found on the three islands. For this reason, it is suggested that the sources of sago palm diversity based on the samples tested are the islands of Papua, Sulawesi and Kalimantan. This data is also consistent with the grouping of sago palm samples through phylogenetic construction at the island level which divides the samples into three groups, namely the Papua group, the Sulawesi group and the Kalimantan group. This indicates that the source of the diversity of sago palm in Indonesia, apart from being on the island of Papua, is also found in other islands, namely Sulawesi and Kalimantan [26].

The genetic relationship of the samples at the individual level shows that the samples are classified into three groups based on the phylogenetic construction. Sago palm relationship studies previously show that sago palm originating from the Malay Archipelago and several samples of sago from Indonesia clustered into two groups and two sub-groups based on the RAPD markers [48]. The sago relationship study focused on the Indonesian archipelago, but with a larger number of samples, showed that the sample individuals were divided into four groups based on RAPD markers [52]. The discrepancy in the division of the number of groups (groups) between the groupings based on cpDNA markers and RAPD markers that was observed in previous studies is something that is often found in studies of genetic relationship using molecular markers. Previous genetic related studies which showed that different molecular markers led to different groupings of certain plants by using cpDNA, RAPD and isozyme markers in Pseudotsuga spp. (Pinaceae) [39], cpDNA and inter-SSR (ISSR) markers in the nucleus genome on *Brassica oleraceae* L. plants using [41], and using cpDNA and mitochondrial DNA on apple plants [24].

Based on cpDNA, the sources of sago diversity in Indonesia are predicted to come from three islands, namely Papua, Sulawesi and Kalimantan. It is suspected that from these three islands, individual sago palm experienced migration in line with migration and population mobilization in Indonesia that had occurred hundreds of years ago. This assumption is reinforced by haplotype data, phylogenetic analysis, and genetic hierarchies which show that samples at the inter-population level and between individuals are significantly different, which means that there are one or more different individuals or populations. Although the source of diversity is found in three islands, if the number of haplotypes is the size of the center of diversity, then the island of Papua is the center of diversity of sago palm in Indonesia because that island is found in the largest number of haplotypes compared to other islands. Apart from the highest number of haplotypes, on the island of Papua, the wild relatives of sago palm

*Genetic Variation*

[27, 28].

in potato have also demonstrated allele polymorphisms in potato [59]. Locus rpl1671, NTCP21, and NTCP22 on sago were detected in three haplotypes out of 10 haplotypes which were specific haplotypes in populations from Jayapura and one specific haplotype each for populations from Serui, Palopo, and Pontianak [26]. The specific haplotype phenomenon is also found in several types of plants i.e. Cunninghamia spp. [60], *Pinus sylvestris* L. [27], and *Alyssum* spp. [29]. The specific haplotypes were found in a population, those indicated the source of diversities in a population. The specific haplotypes of sago palm were found in the populations of Papua, Sulawesi, and Kalimantan indicated the provenance of the diversities, while the most haplotypes of sago palm diverse is the population from Jayapura then followed by the sago palm population from Serui [17]. The large number of haplotypes reflects the high variation in a population in line of the *Abies alba* Mill population [28]. The differences in chloroplast haplotypes in each population reflect differences in genetic entities or sources of variation [29]. The number of haplotypes that were found to be present together in each population is an indication that genetic similarities among individual in a population. It is hypothetically that the sago palm migration by carrying of people. Four haplotypes of 10 haplotypes of sago palm were found in to two or more populations, which means that only four haplotypes were found migration through various kinds of intermediaries. The same thing was also found in *P. sylvestris* L. and *A. alba* Mill. referred to as the refugial population

Population genetic diversity shows that the population from Papua has the highest number of haplotypes (∑H), the number of polymorphic sites (S), and the highest percentage of haplotype polymorphics compared to other populations. A value (HE) equal to one means that no haplotype numbers are the same in individual samples in a population (single haplotype individuals) as happened in the population from Bogor. This is similar with individual haplotype on *P. sylvestris* L [27]. Previous studies on sago palm using RAPD markers showed that sago plant

The genetic hierarchy based on cpDNA was estimated by using analysis of molecular variance (AMOVA) was calculated of differentiation level of population samples at the inter-island level (−3.88% and FCT = −0.03884), between populations within islands (8.49% and FSC = 0.08177), and Papua and others (5.05% and FCT 0.05054) which is low with the probability value not significantly different. High percentage values of variance were observed at the level among individuals (95.39% and FST = 0.04610) and between populations (5.91% and FST = 0.05914) with significantly different probability values [26]. The same thing was also found in *P. sylvestris* L., namely the percentage value of variance between populations (3.24%) with a significantly different probability [27]. The negative value observed at the inter-island level indicates that the sample island level does not contribute to the total measured variance. This phenomenon resembles the tetraploid alfalfa population [47]. Negative correlation coefficients have a biological significance in that the samples at the inter-island level are more closely related than those at the island level [61]. Based on this, it indicates that island or geographic differences do not cause variations in the chloroplast genome, even though the distance between one island and another is far (hundreds to thousands of kilometers). The variation between individuals and between populations contributed 95.39% and 5.91% to the total variety and was significantly different [18]. The results observed were similar with Abies species that was only a small variance value between populations (6.10%), high proportion of variance within the population or between individuals (74.66%) [62]. A low trend of genetic variability between populations is also found in Pinaceae

diversity at the individual level was also high [48, 52].

**36**

[39] and species other than pine [63].

are found. If the data obtained is linked to the incidence of sago distribution in Indonesia, it is strongly suspected that only four haplotypes experienced migration from one population to another, which were then given different local names. The sago population with a specific name for the origin of the Papua region which groups together with populations from other places with other names is also a reflection that in the past these sago populations were only one then experienced joint migration with the migration of people from one island to another or from one population to another. If population migration events have occurred hundreds of years ago and are thought to have caused sago palm to spread from sources of diversity to form new populations or join old populations on islands that are sources of diversity, it is still possible because the measure of similarity is cpDNA, which has very conservative sequences [64], a very low mutation rate of between 3.2 x 10–5 and 7.9 x 10–5 [23], is not recombinant [24, 64] and are inherited uniparental [25, 39].

#### **3.4 Genetic improvement by using marker-assisted breeding (MAB)**

The development of genetics and technology molecular has facilitated our understanding of the genetics underlying the traits sought by plant breeding. The development of molecular markers allows plant breeding to develop faster and more advanced in producing superior organisms. The benefits of DNA markers are for germplasm characterization, selection of desired traits from genomic regions involved in the expression of traits of interest, and single gene transfer. The application of selection using efficient and effective markers to improve polygenic properties certainly requires new technology. Genetic improvement of sago palm may use transformation agrobacterium-mediated and particle bombardment. Successfully introgression *bar* and *gus* gene into sago palm genome [65]. The embryogenic callus was the most appropriate transformation material compared to the via callus, the embryoid stage and the shoots initiated by using *Agrobacterium*mediated. The transformation of the gene gun demonstrated greater efficiency of transformation than those transformed with Agrobacterium when targets were bombarded once or twice with 280 psi helium pressure at a distance of 6 to 8 cm [65]. Therefore, economics interesting genes may introgression into sago palm genome in the future.

The purpose of MAB is to enhance certain characteristics in plant or animal breeding programs. Strategy for rapidly integrating a targeted gene into a wheat genotype in only two generations and restoring 97% or more of the recurrent genotype of the parent by using MAB [66]. Deconvolution of ancestry offers a first step towards selection of suitable admixture profiles at the seed or seedling level, which will support marker-assisted breeding aimed at introgressing wild Vitis species while maintaining the desirable characteristics of elite *V. vinifera* cultivars [67]. Marker-assisted backcrossing can be used in plant breeding to integrate traits into elite cultivars while minimizing the transfer of unwanted alleles from the donor genome [68]. This method includes the selection of foreground as well as context. Foreground selection refers to offspring screening and selection based on the presence or absence of a particular allele associated with a feature of interest. Conversely, selection of offspring on the basis of genomic ancestry estimates is the history selection.

The MAB needs to be developed to accelerate and increase the success of the breeders to produce superior seeds. Recently, breeders were developed abundant MAB linked with specific characters of plant genetics. Simple sequence repeat (SSR), namely Md-PG1SSR10kd tightly linked with fruits texture of apple [69] and microsatellites RM5926 and AP5659–5 were developed for detecting rice

**39**

**Author details**

Barahima Abbas

Manokwari, Indonesia

*Adaptive Evolution and Addressing the Relevance for Genetic Improvement of Sago Palm…*

blast resistance genes, those markers tightly linked with *Pi*-1 and *Piz-*5 genes respectively [70]. Marker-Assisted Introgression of b-carotene hydroxylase was developed for detecting b-Carotene Rich in maize hybrid [71]. Furthermore, Muthusamy et al. (2014) stated that B-carotene concentration among crtRB1 introgressed inbred ranged from 8.6 to 17.5 mg/g - a maximum increase up to 12.6 times over recurrent parent. In comparison to 2.6 mg/g in the original hybrid, the reconstituted hybrids formed from improved parental inbred also showed enhanced kernel *b-carotene* as high as 21.7 mg/g [71]. This study may use as a model for increasing quality starch that resulting of sago palm and other plant in the

Genetic resources of sago palm in Indonesia were explicated as follows: (1) Characters of sago palm in Indonesia were shown varied based on cpDNA markers and large variation based on RAPD markers. (2) Variation of starch production of sago palm correlated with *Wx* genes variation, (3) Distances barrier and geographies isolation in line of sago palm dispersions in Indonesia (4) Characteristics of genetic were observed does not related with vernacular names those were given by local people (5) Papua islands, Indonesia territorial is proposed the center of sago palm diversities, (6) Papua islands, Sulawesi islands and Kalimantan islands will be the provenance of the diversities (7) Genetic improvement of sago palm might enhanced using molecular marker that link to interesting genes by developing

This work is supported by Research Development Project No.198/SP2H/LT/ DRPM/2020 from the Directorate General of Strengthen Research and Community

Faculty of Agriculture, Post Graduate Program, University of Papua (UNIPA),

© 2020 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

\*Address all correspondence to: b.abbas@unipa.ac.id

provided the original work is properly cited.

Service (DRPM-DIKTI), Republic of Indonesia.

*DOI: http://dx.doi.org/10.5772/intechopen.94395*

current time and in future time.

**4. Conclusions**

marker-assisted breeding.

**Acknowledgements**

*Adaptive Evolution and Addressing the Relevance for Genetic Improvement of Sago Palm… DOI: http://dx.doi.org/10.5772/intechopen.94395*

blast resistance genes, those markers tightly linked with *Pi*-1 and *Piz-*5 genes respectively [70]. Marker-Assisted Introgression of b-carotene hydroxylase was developed for detecting b-Carotene Rich in maize hybrid [71]. Furthermore, Muthusamy et al. (2014) stated that B-carotene concentration among crtRB1 introgressed inbred ranged from 8.6 to 17.5 mg/g - a maximum increase up to 12.6 times over recurrent parent. In comparison to 2.6 mg/g in the original hybrid, the reconstituted hybrids formed from improved parental inbred also showed enhanced kernel *b-carotene* as high as 21.7 mg/g [71]. This study may use as a model for increasing quality starch that resulting of sago palm and other plant in the current time and in future time.
