Amorphophallus is a perennial herb of Araceae. According to statistics, more than 200 species of Amorphophallus have been found all over the world. There are 26 species in China and 14 of them are endemic to China. They are widely distributed in China's Yunnan, Sichuan, Guizhou, Chongqing and western Hubei and other shady areas. Amorphophallus is the only cash crop in the world that can produce a large amount of KGM (Liu, 2004, China Agricultural Press, pp.1-84). KGM is an edible dietary fiber, which has significant effects on weight loss, detoxification, blood lipid and blood glucose, etc. (Xie et al., 2005). In recent years, People's pursuit of health has set off an upsurge of Amorphophallus processed food. The research reports on Amorphophallus related food and materials have been in-depth, but the research reports of Amorphophallus molecular biology have just started and gradually increased in recent years. Therefore, this paper summarized the research progress of Amorphophallus in karyotype, molecular marker application, omics research and functional genes in recent years, hoping to provide a reference for Amorpho-phallus

1 Karyotype of Amorphophallus

Karyotype generally refers to the type of chromosome. It is a basic method for studying chromosomes, and it is also an important means to understand biological genetic variation, the evolution of biological systems, and the genetic relationship between species. The karyotypes of Amorphophallus chromosomes are 2n=24, 2n=26, 2n=28 and 3n=39, among which the 2n=26 karyotype is the most common in Amorphophallus. Liu et al. (1985) compared Amorphophallus konjac and Amorphophallus albus and found that they both were diploid (2n=26), and their karyotypes were 2n=16m+2Sm+8st (2SAT) and 2n=16m+6Sm (2SAT)+4ST. Zheng and Liu (1989) compared the karyotype and Giemsa banding analysis of Amorphophallus konjac, Amorphophallus albus, Amorphophallus dunnii, Amorphophallus mairei, and the chromosomes of the four species were all 2n= 2x=26, both are 2B karyotypes. The degree of evolution from high to low is Amorphophallus konjac, Amorphophallus dunnii, Amorphophallus mairei, Amorphophallus albus. Wei and Wen (2003) analyzed the number and karyotype of Amorphophallus coaetaneus. It shows that the chromosome number of Amorphophallus coaetaneus is 2n=26, and the karyotype formula is 2n=2x=26=20 m+6 sm. It belongs to the "2B" type. Compared with other species of the Amorphophallus, Amorphophallus coaetaneus is more primitive. Niu et al. (2008) applied colchicine to the diploid of Amorphophallus albus. The induced autotetraploid Amorphophallus albus were 2B karyotype. Zhou et al. (2011) first conducted a karyotype analysis on the chromosomes of Amorphophallus bulbifer root tip cells. The results showed that the number of somatic chromosomes of Amorphophallus bulbifer was 2n=39, and the karyotype formula was: 2n=3X=39=33 m+ 6 sm, is the most evolved Amorphophallus species in this genus. Zhang (2014) further analyzed the karyotype of Amorphophallus. The number of chromosomes has four karyotypes: 2n=24, 2n=26, 2n=28 and 3n=39. The chromosome types are as follows: 2A, 1B, 2B, 3B. From the report of Amorphophallus karyotype, we can find that the genetic variation of Amorphophallus is very rich.There are not only differences in karyotype chromosomes between different species, but also karyotype differences between the same species. These complex genetic backgrounds indirectly affect the progress of Amorphophallus molecular research.

2 Application of Molecular Markers

Compared with traditional genetic markers such as morphological markers, cytological markers, and biochemical markers, molecular markers are genetic markers for detecting nucleic acid polymorphisms, which directly reflect genetic variation at the DNA level. Because of its advantages such as large number of markers, high polymorphism, wide distribution, neutrality, and no environmental restrictions, it is widely used in various research and practice (Gavin et al., 1994). The development of PCR technology has led to the explosive development of new technologies in molecular biology. However, only a handful of these techniques, namely RFLPs, RAPDs, AFLPs, ISSRs, SSRs and SNPs have received global acceptance (Grover and Sharma, 2016).

Zhang and Sun (2006) first used RAPD molecular marker technology to PCR amplify the genomic DNA of Amorphophallus albus. The cluster analysis results are divided into 5 categories. Comparative analysis found that the classification is not closely related to the distribution of petiole markings, but has obvious correlation with the region. This result provides molecular evidence for the genetic relationship within the Amorphophallus albus. Teng et al. (2008) conducted DNA polymorphisms on the germplasm of different kinds of Amorphophallus. This study used ISSR technology to identify wild sample varieties, and used AFLP marker technology to prove that Amorphophallus resources in different regions have rich genetic diversity. This is the first application of ISSR and AFLP markers in Amorphophallus research, laying a theoretical foundation for research on molecular genetics and breeding of Amorphophallus. Chen (2010) improved the traditional DNA extraction method of Amorphophallus, and used two molecular markers RAPD and ISSR to analyze the regenerated Amorphophallus from petiole callus, revealing the somatic clonal variation of its regenerated plants. Zheng et al. (2013) first attempted to develop a large number of SSR markers for Amorphophallus konjac and Amorphophallus bulbifer based on the transcriptome database. Using validated primers, the diversity of 25 plants was analyzed. The establishment of SSR markers in the institute is a valuable resource for studying the genetic diversity, linkage map and germplasm characteristics of Amorphophallus (Araceae). Ren and Pan (2013) used ISSR molecular markers to analyze the population genetic diversity of five Amorphophallus species in southern Yunnan, China. This study is not sufficient as evidence to distinguish species and Interspecific, but it can provide a certain basis for the division of Amorphophallus species and the judgment of the relationship interspecific. Zhang (2014) analyzed and clustered Amorphophallus varieties using ISSR technology, and verified that the clustering results are consistent with the cytological classification results based on the number of chromosomes. Pan et al. (2015) selected some individuals from 10 wild populations in Chinese middle-area and used AFLP technology to obtain the results of high genetic variation in the Amorphophallus. Mekkerdchoo et al. (2013) used RAPD markers to track the evolution of KGM in Araceae plants, and screened a high-economic variety of Amorphophallus. Gholave et al. (2017) used RAPD and ISSR techniques to classify Amorphophallus in order to reconstruct the relative species of Amorphophallus. In this study, a development method of Amorphophallus SSR based on restriction-site-associated DNA (RAD-seq) was reported. Yin et al. (2020) used restriction-site-associated DNA (RAD-seq) to establish SNP markers in order to understand the genetic diversity and population structure of the endangered species Amorphophallus krausei, laying the foundation for the conservation genetics of Amorpho-phallus krausei.

At present, the main molecular markers used in Amorphophallus are random DNA molecular markers (RDMs) and functional molecular markers (FMs), including five markers: RAPD, AFLP, ISSR, SSR, and SNP. These techniques are mainly used for research on genetic diversity of Amorphophallus, classification and identification of species, varieties and hybrids, and genetic relationship, but there are few reports in the research of Amorphophallus genetic breeding and phylogenetic analysis.

3 Omics Research of Amorphophallus

3.1 Genomics

The genome is the sum of all the genetic material of an organism. It contains coding DNA and non-coding DNA, mitochondrial DNA and chloroplast DNA. By systematically studying the genome sequence of a certain plant, the sequence information of the plant's genome and important functional genes can be obtained, which provides an important reference for studying the structure, composition, function and evolutionary laws of this plant. Because of the huge genome-wide data of Amorphophallus, there are many differences in karyotypes among species, and there are large genetic variations within species. These have brought great difficulty to the sequencing work, so there is currently no whole genome sequencing data of Amorphophallus.

Compared with the whole genome, the Amorphophallus chloroplast genome data is relatively small, about 150 kb. Liu et al. (2019) used high-throughput Illumina sequencing technology to sequence and assemble the Amorphophallus chloroplast genome. The results indicate that the complete Amorphophallus chloroplast gene consists of large fragments and small single-copy regions, separated by a pair of inverted repeats, and has 113 unique genes (80 protein-coding genes, 29 tRNAs and 4 rRNAs). Phylogenetic analysis shows that Amorphophallus is closely related to taro and pinellia (Hu et al., 2019). The study of complete chloroplast genome information can provide new research directions for the commercial breeding and medicinal development of Amorphophallus.

3.2 Transcriptomics

Transcriptome analysis is of great significance for predicting certain gene functions in the growth, development and metabolism of Amorphophallus. Zheng et al. (2013) sequenced Amorphophallus transcriptome and identified 10,754 SSR markers, and successfully verified 77 polymorphic markers in 25 individuals. These research results provide a theoretical basis for studying the genetic diversity and germplasm characteristics of Amorphophallus. Diao et al. (2014) constructed two transcriptomes of Amorphophallus konjac and Amorphophallus bulbifer. All 108,651 unigenes with average lengths of 430 nt in Amorphophallus konjac and 119,678 unigenes with average lengths of 439 nt were generated from 54,986,020 reads and 52,334,098 reads after filtering and assembly, respectively. Through comparison with the Nr, Swiss Prot, KEGG and COG databases, a total of 80,332 transcripts are differentially expressed between Amorphophallus konjac and Amorphophallus bulbifer. This research promotes the study of molecular genetics of Amorphophallus. Zhong et al. (2018) found possible sphingolipid metabolism pathways and 13 enzymes involved in the transcriptome analysis of Amorphophallus muelleri, and studied the key enzyme neutral ceramidase, proving that Amorphophallus muelleri has great potential in anti-virus. Li et al. (2019) compared Amorphophallus albus and Amorphophallus konjac with Arabidopsis thaliana by tBlastn comparison and splicing. It is inferred that the GDP-mannose pyrophosphorylase gene GMP in Amorphophallus is full-length, which further proves that the GMP gene is very likely to be related to stress resistance. Li et al. (2020) analyzed the transcriptome data of the leaf buds and flower buds of Amorphophallus konjac and found that 35 genes are involved in the plant hormone signaling pathway and hormones play an important role in the differentiation of leaf buds and flower buds.

3.3 Proteomics, metabolomics, and miRNAs

The proteomics and metabolomics are of great significance to the development of Amorphophallus molecular biology research. However, there are no relevant reports on the application of proteomics and metabolomics in the research of Amorphophallus. There are few research reports on miRNAs (MicroRNAs). Only Diao et al. (2014) sequenced and obtained 5,499, 903 small RNA sequences in Amorphophallus leaves, and predicted potential targets of miRNAs based on transcripts.

This research described KGM biosynthesis genes involved in the leaves. It is helpful to further understand its key role in carbohydrate synthesis and other important metabolic pathways in Amorphophallus. The next research can analyze the quantitative description of the target protein expression and the expression changes under the influence of other factors, the quantitative analysis of certain protein expression data of Amorphophallus, combined with metabolomics, so as to clarify its gene expression regulation. This provides new ideas for the development of research on the growth and development of Amorphophallus. 

4 Functional Genes Research of Amorphophallus

4.1 Genes related to KGM synthesis

Amorphophallus is currently the only plant capable of producing large amounts of KGM in its body. KGM is the main biologically active component present in Amorphophallus bulbs. Many colorless and bright particles with a diameter of about 1 mm can be clearly seen by cutting the bulb tissue. These particles are called glucomannan cell. The deposition of glucomannan in glucomannan cells follows time regulation, and its expression increases at the end of the vegetative growth cycle (Chua et al., 2013). Gille et al. (2011) proposed the nucleotide sugar interconversion pathway from sucrose to GDP-mannose and GDP-glucose, the two building blocks of glucomannan as well as the starch biosynthesis pathway. The highest number of ESTs for this pathway was found to be from CSLA3—a putative glucomannan synthase (Figure 1).

Figure 1 glucomannan and starch biosynthesis pathway in the Amorphophallus konjac Note: The necessary converting enzymes found in the EST data are given in red with the number of obtained reads in green. The thickness of the arrows is representative for the number of reads. Reference (Gille et al., 2011)

The first gene related to glucomannan synthesis found and identified in plants is galactosyltransferase, which can cause the mannan backbone to be replaced and make the polymer more soluble (Edwards et al., 1999). The glucomannan in the cell wall of the model plant Arabidopsis thaliana is synthesized by members of the cellulose synthase-like family A (CSLA) (Goubet et al., 2009). The polymer is synthesized from activated nucleotide sugars-GDP-mannose and GDP-glucose (Liepman et al., 2005). Gille et al. (2011) found that high levels of cellulose synthase-like family A (CSLA) transcript AkcsKa3 is a glucomannan synthase produced in Amorphophallus bulbs. The glucomannan synthase uses GDP-D-mannose and GDP-D-glucose as substrates to synthesize storage glucomannan in the bulb. Li et al. (2019) speculated that the full-length sequence of the GDP-mannose pyrophosphorylase (GMPase) registered by Gille et al. in GeneBank has certain problems. They re-sequenced the full-length sequence of Amorphophallus GMPase and cloned the promoter sequence of the gene to verify their conjecture. Zhang (2018) analyzed the expression of starch synthesis gene AmAGP in Amorphophallus muelleri. As the bulb expands, the expression of AmAGP is negatively correlated with the actual KGM content. Studying the structure and function of AmAGP of Amorphophallus will help improve the quality of Amorphophallus muelleri and increase the content of glucomannan. Shi et al. (2020) cloned ADP-glucose pyrophosphorylase (AGP) from Amorphophallus muelleri and further verified that AmAGP is involved in the regulation of starch synthesis.

At present, the biosynthetic pathways of starch and glucomannan in Amorphophallus bulbs have been proved, but the specific regulation relationship between the two at the molecular level is still unclear, and further research is needed.

4.2 Genes related to soft rot resistance

In recent years, Amorphophallus industry has developed rapidly, and the problems of plant diseases and insect pests in production have received extensive attention from researchers. Among them, soft rot is the most prominent. Soft rot is a bacterial disease. The pathogens that cause soft rot are mainly E.carotovora subsp. carotovora (Ecc), E.chrysanthemi (Ech) and E. carotovora subsp. Atroseptica (Eca) three pathogens and other unidentified species (Wu et al., 2010).

In view of the difficult control soft rot disease, the screening of disease resistance genes of Amorphophallus is particularly important. Plant disease resistance genes (R genes) are a type of genes in the plant genome that can induce R protein to resist infection and expansion of pathogenic bacteria and play an important role in the genetic regulation mechanism of plant disease resistance (Nimchuk et al., 2001). At present, it has been found that the amino acid sequences of the cloned plant disease resistance genes have high homology, and they all show specific conservative structures, such as nucleotide binding site (NBS), leucine-rich repeats (LRR), leucine zipper (LZ) and so on. Lei et al. (2019) designed primers based on the conservative regions of the known plant NBS-LRR resistance gene analog, RGA. They cloned a homologous fragment of the soft rot resistance gene from the disease-resistant plant of the Amorphophallus konjac to obtain a complete disease-resistant R gene.

Wei et al. (2019) screened and synthesized inverted primers from the reported NBS-LRR-like conserved domains of plant disease resistance genes. They amplified and isolated 4 disease-resistant gene homologous sequences with high homology to known disease-resistant genes from Amorphophallus konjac. The accession numbers in NCBI database are MK123507, MK123508, MK036024, MK036028. The current research on the disease resistance genes of Amorphophallus only has homologous sequences of NBS-LRR disease resistance genes, and no one has screened and cloned other disease resistance R genes. This provides a reference for the next development of new disease-resistant varieties and research on disease-resistant transgenic breeding.

5 Discussion and Future Objectives

At present, although there have been many research reports on molecular biology of Amorphophallus, the system has not yet formed. And it needs further in-depth and broad research. In the omics research, there are no whole genome sequencing reports for all plants of Amorphophallus and Araceae. Because the Amorphophallus genome data is too large, this further increases the difficulty of genome sequencing. The lack of genome-wide data will affect the analysis of transcriptome and proteome data, and also limit the development of molecular markers. Compared with genomics, transcriptome research has been initially applied in Amorphophallus, involving the differentiation and stress resistance of flowers and leaves. Transcriptome sequencing also provides conditions for the large-scale development of SSR markers, which makes it possible to quickly identify a large number of SNPs in the genome. There are no reports on proteomics and metabolomics, and there are few research reports on miRNAs. In the application of molecular markers, it is only used to distinguish the types or varieties of Amorphophallus, without forming a comprehensive system. In functional gene research, the main focus is on regulating the formation of KGM and research on NBS-LRR disease resistance genes. There is little exploration of other genes.

In the future, related research on molecular biology of Amorphophallus should mainly focus on the following aspects: First, carry out the sequencing of Amorphophallus genome, combining transcriptome and proteome data, and multi-omics joint analysis. Omics research is a good method to discover differential genes and lay the foundation for further identification of gene functions. The second is to combine Amorphophallus genome and transcriptome data to obtain molecular markers for important economic traits. Through molecular marker-assisted breeding, combining traditional breeding and biotechnology breeding to breed more valuable varieties with broader market prospects. The third is to conduct a more comprehensive study in functional genes of Amorphophallus, such as the functional identification and regulation of genes in flower morphogenesis, bead formation, stress resistance, growth and metabolic pathways.

Acknowledgements

The funding for this study was jointly funded by the National Natural Science Foundation of China (31760349) and (31960071).

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