Anther Culture in Crop Plants: Progress and Perspectives

M. Thoihidul Islam; Mohammad Rashid Arif; Md. Toufiq Hasan; Arif Hasan Khan Robin

doi:10.9787/PBB.2023.11.2.69

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Anther Culture in Crop Plants: Progress and Perspectives

M. Thoihidul Islam¹, Mohammad Rashid Arif¹, Md. Toufiq Hasan², Arif Hasan Khan Robin^1,*

Plant Breeding and Biotechnology 2023;11(2):69-96.
Published online: June 1, 2023

DOI: https://doi.org/10.9787/PBB.2023.11.2.69

¹Department of Genetics and Plant Breeding, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh

²Faculty of Agriculture, Bangladesh Agricultural University, Mymensingh 2202, Bangladesh

*Corresponding author Arif Hasan Khan Robin, gpb21bau@bau.edu.bd, Tel: +88-09167401-7/64714, Fax: +88-09161510

• Received: October 11, 2022 • Revised: February 24, 2023 • Accepted: March 8, 2023

This is an open-access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0) which permits unrestricted non-commercial use, distribution, and reproduction in any medium, provided the original work is properly cited.

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Abstract
INTRODUCTION
CONCLUSION
References

Abstract

A resurrection has started in haploid and double haploid research in the twenty-first century. The haploid and double haploid could be achieved through in vivo and in vitro anther and microspore culture techniques. Fixing the homozygosity is the most striking benefit of androgenesis. Various factors like genotypic dependency, growth condition, developmental stage of the microspore, pre-treatment, culture media, regeneration media, growth hormones, and various chemicals have a direct effect. Wheat, rice, Brassica, and tobacco are the notable crops where anther and microspore culture has been utilized. These haploidy and double haploidy through anther culture served many purposes of basic and applied research. Especially, double haploid cultivars have been cultivating around the globe. In addition, for chromosome mapping, QTL mapping, marker-assisted selection, marker-assisted backcrossing, mutation breeding, genome-wide association study, genomic engineering, and genome editing, androgenesis based haploid and double haploid plants have been exploited due to the effectiveness. Recently, researchers are trying to explain albinism that happens during anther culture from an epigenetic perspective. Further prospects of haploid and doubled haploid research through anther culture have been described in this review.
Key words: Haploid, Anther culture, Microspore culture, Rice, Wheat, Brassica, Tobacco, Albinism

INTRODUCTION

Haploid plants are those which contain gametophytic chromosome number and it indicates plants contain a single set of chromosome (Germanà 2011). When the haploid plants undergo spontaneous or induced chromosome dou-bling procedures like- colchicine treatment, nitric oxide, or oryzalin treatment they become double haploid or fertile homozygous diploid (Mishra et al. 2016). Though the haploid plants are sexually sterile, the double haploids are fertile and pure homozygous lines (Devaux and Pickering 2005). The haploid and double haploid plants could be obtained through in vivo and in vitro techniques.

In vivo methods includes field condition-based or com-bination of field and control condition based techniques like parthenogenesis (Lacadena 1974; Chase 1969; AI- Yasiri and Rogers 1971; Dumas deVaulx and Pochard 1974; Khush and Virmani 1996), semigamy (Dunwell 2010; Turcotte and Feaster 1963, 1969; Zhang and Stewart 2004), twin seedlings (Carl and Yawney 1972; Nezhevenko and Shumnyi 1970), chromosome elimination (Khush and Virmani 1996; Kasha and Kao 1970; Jensen 1977, 1983; Pickering 1980; Devaux 2003; Furusho et al. 1991), crossing between different ploidy level (Dunwell 2010; Asker 1983; Oiyama and Kobayashi 1993; Germanà and Chiancone 2001; Murovec and Bohanec 2011; Bosemark 1971). By contrast, the in vitro techniques are control condi-tion based. Gynogenesis and androgenesis are the in vitro methods of gametic embryogenesis. Gynogenesis is the haploid plant regeneration procedure where the haploid embryos are produced from in vitro culture of female game-tophytes. The success of this method is greatly affected by the genotype of the parent and environment (Murovec and Bohanec 2011). These limiting factors turn this method into less efficient and gynogenesis only utilizes when other efficient methods have a lower response (Forster et al. 2007).

Androgenesis is the process of haploid production through the culturing of anther or microspores in vitro conditions (Fig. 1). Anther culture is a widely utilized technique for developing haploid and double haploid plants because of its simplicity and effectiveness. It covers a vast range of genotypes with higher plant establishment (Sopory and Munshi 1996). Also, haploid plant regeneration is possible through isolated microspore culture. Due to its delicacy, this technique demands sound technique and improved equipment compared to anther culture (Germanà 2011).

In vitro anther culture is an empirical study where crops like wheat (Triticum aestivum L.), barley (Hordeum vulgare L.), tobacco (Nicotiana spp.), rapeseed (Brassica napus L.) showed high regeneration capability. This specialty made these crops model species to study the underlying mechanism (Forster et al. 2007). The gametic embryogenesis to develop haploid plants had been studied by plant scientists for above around 5 decades and the notable review articles are Magoon and Khanna (1963), Kasha (1974), Zhang et al. (1990), Jain et al. (1996–1997), Smykal (2000), Maluszynski et al. (2003a, b), Andersen (2005), Palmer et al. (2005), Xu et al. (2007), Seguı`- Simarro and Nuez (2008a), Touraev et al. (2009), Dunwell (2010), Seguı`-Simarro (2010), Germanà (1997, 2006, 2007, 2009, 2011). Then, Maluszynski et al. (2003a) stated fourty four protocols of developing double haploid from around thirty-three plant species.

This current review will try to address the history and crop-wise (rice, wheat, Brassica, and tobacco) success and utilization status of androgenesis along with albinism which is termed as the major drawback.

Importance of haploid and double haploid

The plant researchers have predicted the potentiality of haploid and double haploid (DH) in plant science research which was established in further years (Sunderland 1978; Forster and Thomas 2005). The pure line is mandatory for developing a hybrid variety and developing a pure line requires several generations by conventional approach (Keles et al. 2015). Haploid and DH just cut off the duration and the homozygous pure line developed within a single generation and it is a major breakthrough in cultivar development (Datta 2005; Dunwell 2010). Afterward, in vegetable breeding, various parental lines show “inbreeding depression” and they only outyield in F1 condition after crossing with another parent (Maluszynski et al. 2001; Hochholdinger and Hoecker 2007). The hybrid perfor-mance could be fixed in a single line by using DH techno-logy (Germanà 2011). Crops like rye (Secale cereale) (Immonen and Anttila 1996) and forage grasses (Nitzche 1970) are recalcitrant to develop fertile homozygous by selfing. DH development of those crops is the best app-roach to confront the recalcitrance (Germanà 2011).

Marker-assisted breeding along with the DH technique has been successfully utilized in the various breeding programs to cut short the breeding cycle (Wessels and Botes 2014; Tuvesson et al. 2007; Mago et al. 2011). Then, the marker-assisted backcross breeding DH approach was integrated to reduce the duration and the target trait was introgressed in an elite line within the quickest time (Toojinda et al. 1998).

The haploid and DH are also important tools in mutation breeding. The homozygous mutant line was developed by inducing mutation in the single gametic cells which gone through embryogenesis and double haploid mutant plant population achieved (Szarejko and Forster 2006). Alterna-tively, the mutation was used in the gamete of the M1 plant to get the DH plant. In barley, rice, and wheat it has been utilized successfully (Forster et al. 2007). Noteworthy, the unicellular microspore is recommended for mutation induction (Germanà 2011). Then, the trait developed by mutation is recessive, and sometimes it is found in M2 or M3 generations. In that case, double haploidy is a good option; it just fixes the desired recessive trait (Germanà 2011; Szarejko and Forster 2007). TILLING (Targeting Induced Local Lesions In Genomes; McCallum et al. 2000; Perry et al. 2003) is a reverse genetic approach which applied to relate the gene with the phenotypes by using chemical mutation along with the high throughput screen-ing method like SNPs (single nucleotide polymorphisms). Due to the inherent variation, a false-positive result could be achieved from the starting material (Germanà 2011). To resolve this problem, in vitro methods like anther or microspore culture is a good alternative (Tadele et al. 2010).

Double haploid and haploid have strengthened its position in applied research and basic research of breeding and genomics. The DH lines have homozygosity and uniformity; moreover, they can be replicated multiple times in various locations. This facility makes double haploid ideal for genetic mapping (Tinker et al. 1996; Khush and Virmani 1996). Likewise, in genetic and physical mapping the double haploid has considerable precision to locate the targeted gene (Kunzel et al. 2000; Wang et al. 2001). The chromosome map of various crops including barley, rice, rapeseed, and wheat had constructed using the haploid and double haploid plant (Forster and Thomas 2005). In the ‘gametoclonal variation’ variation study the haploids have great potentiality as it points out the morphological, biochemical, and chromosomal variation within the plants derived through gametic cell culture (Evans et al. 1984; Morrison and Evans 1987). Also, the variations due to segregation, independent assortment, chromosome doubling procedure, and diploid level are explained from the ‘gametoclonal variation’ (Morrison and Evans 1987; Huang 1996).

Potrykus and his co-authors predicted the potentiality of the anther in genetic transformation as anther cultured isolated pollen has remarkable regeneration frequency (Potrykus et al. 1985; Potrykus 1988) which has proved further. In genetic transformation, the single-celled micro-spore, haploid embryo, or calli act as the recipient of the gene of interest. Thus, the obtained DH plant will contain the transgene in the homozygous state. Microinjection, electroporation, particle bombardment, and Agrobacterium tumefaciens-mediated transformation were the gene deli-very technique was found effective in microspore based gene delivery (Touraev et al. 2001; Chen et al. 2006; Chauhan and Khurana 2011; Li et al. 2012; Ohnoutková and Vlčko 2020).

Genome editing is the state-of-the-art technology of the twenty-first century where transgene is not required to achieve genetic variation. Zinc Finger Nucleases (ZFNs), Transcription Activator-Like Effector Nucleases (TALENs), and Clustered Regularly Interspaced Short Palindromic Repeats and CRISPR-Associated Protein 9 (CRISPR/ Cas9) are the popular genome editing techniques. Among those, CRISPR/Cas9 is the most updated and efficient tool to edit any organism. Generally, immature embryos are used as the ex-plant of transformation. By contrast, studies show that anther or microspore has feasibility over the conventional method because the availability of gene-tically and physiologically identical microspore or anther is relatively easy. Then, after diploidization (spontaneous/ induced) the plants are in a homozygous state whether it might take several generations to achieve the homozy-gosity. Afterward, due to the haploidy status, the gene to be edited will be half in number which increases the efficacy (Bhowmik et al. 2018; Han et al. 2021). In essence, microspore or anther culture-based haploid transformation will save time, space, and money with great effectivity (Bhowmik et al. 2018; Han et al. 2021; Bilichak et al. 2020; Haque et al. 2018; Brandt et al. 2020; Borisjuk et al. 2019; Jansing 2019).

A brief history of success of anther culture in haploid and doubled haploid plant regeneration in crop plants

The haploidy status of the plant was first observed in sporophytic weed species Datura stramonium (Jimson weed) in 1921 with proper cytological evidence (Blakeslee et al. 1922). In another report, Harland, 1920 claimed about the haploid presence in a sea island cotton commonly named as “Man cotton” (Harland 1920, 1936, 1955). Then, the occurrence of the first androgenic haploid was stated by Kostoff (1929) found in tobacco. That contemporary times, the concept of pure lines and gene theory proposed by Wilhelm Johannsen in 1903 and 1909 respectively, also the phenomenon of induced mutation by H.J. Muller’s (1927) published. These concepts helped to fathom the poten-tiality of haploids and double haploids in genetics and plant breeding, as a result, haploid and double haploid study gained the center of interest in further decades (Maheshwari 1996). From 1920 to 1950, plant scientists intensively tried to develop haploid plants but at that time anther culture was not a familiar tool to researchers (Maheshwari 1996). Finally, in 1964, Guha and Maheshwari generated haploid embryos through in vitro anther culture from the immature anther of Datura innoxia. This novel discovery coincided with the prediction of Katayama and Nei (1964) where they stated the potentiality of pollens in generating haploid plants. This discovery helps to initiate extensive research in Poaceae, Brassicaceae, and Solanaceae family (Germena 2011). Subsequently, a similar result was reproduced for tobacco in France and Japan (Bourgin and Nitsch 1967; Nitsch and Nitsch 1969; Nakata and Tanaka 1968). Then, Niizeki and Oono (1968) successfully generated an andro-genic haploid rice plant which is a major cereal crop globally. These breakthrough findings raised a pinnacle of interest in haploids so that they could achieve homozygous pure-line within a single generation (Nitsch and Noreel 1973; Nitsch 1974a, b). As interest rises, the first inter-national symposium was held at the University of Guelph, Canada with the title of ‘Haploids in Higher Plants’ in 1974 (Kasha 1974).

The first double haploid crop variety was “Maris Haplona” of rapeseed mustard (Brassica napus) released at the begin-ning of the 1970s (Thompson 1972). Afterward, a Barley (Hordeum vulgare) cultivar named Mingo released in 1980 which was also a DH crop variety (Ho and Jones 1980). Noteworthy, the early twenty-first century has experienced a resurrection in haploid research especially in higher plants (Forster et al. 2007). Many initiatives had been taken to add value to this workforce. European Union ran a project COST 851 program (https://www.cost.eu/actions/851/#tabs|Name:overview) where a research network de-veloped to work on the topic “Gametic cells and molecular breeding for crop improvement” for six years (Germena 2011). Significantly around 300 DH crop varieties have developed using various double haploid inducing techni-ques including in vitro anther culture (Germanà 2011), and around 200 species regenerated from androgenesis of Gramineae, Solanaceae, and Cruciferae family to date (Germanà 2011; Dunwell 1986; Hu and Yang 1986).

Rice perspectives

Rice is one of the principal crops, feeding more than half of the world population (http://ricepedia.org/rice-as-food/the-global-staple-rice-consumers). The first haploid plant of rice was developed in 1968 (Niizeki and Oono 1968). Considering the efficiency and regeneration potential androgenesis especially anther culture is the simplest and efficient technique in cereal crops including rice (Forster et al. 2007; Murovec and Bohanec 2012; Mishra and Rao 2016). Remarkably, around 50-60% of double haploid plants regenerate automatically along with haploid plants in rice anther culture and it is a special hallmark for rice anther culture (Maluszynski et al. 2003a, b).

Sarao and Gosal (2018) and Mishra and Rao (2016) discussed thoroughly in their reviews regarding rice androgenesis. In rice anther culture, the genotype is the most deciding factor (Mishra and Rao 2016). Among species, indica has a very limited response to anther culture (Chen et al. 1991). By contrast, japonica species have higher regeneration efficiency (He et al. 2006). Then, early or mid- to late-uninucleate are the best stages for haploid regeneration (Datta and Wenzel 1998). Pretreatment of rice pollen is essential where various stresses like temperature shock, osmotic shock, sugar starvation were implied to a various extent (Mishra and Rao 2016; Datta 2001). Even though, gamma irradiation had utilized to treat anther which proved as a useful technique (Zapata and Aldemita 1989; Chen et al. 2001). Afterward, colchicine was used other than a mutagenic agent to promote gametic embryo-genesis in rice (Alemano and Guiderdoni 1994).

As culture media, N6 media with a trace amount of NH⁴⁺ is widely used in rice anther culture. Especially for indica species, Raina and Zapata (1997) recommended MO19 medium. The culture media needs carbohydrate source. Sucrose is commonly utilized for these purposes but maltose has supremacy in a few contexts compared to sucrose, such as albino plant frequency reduced by maltose utilization (Reinert and Bajaj 1977; Sen et al. 2011; Park et al. 2013). Besides these - sorbitol, amino acid, and AgNO³ were found beneficial for plant regeneration from andro-genesis in rice (Yoshida et al. 1994; Ogawa et al. 1995; Faruq et al. 2014). Growth regulators are also used in the in vitro culture of anther and it has a positive impact on in vitro culture (Mandal and Gupta 1995). The most com-monly utilized growth regulator for callus induction from rice anther is 2,4-dophenoxy acetic acid (2,4-D) and naphthalene acetic acid (NAA) where both of them are auxin (Trejo-Tapia et al. 2002). But for direct androgenesis Indole acetic acid (IAA) and NAA might be used (Ball et al. 1993). In special cases, growth regulators like kinetin and benzyl amino purine are recommended where 2,4-D or NAA cannot serve properly (Mandal and Gupta 1995). Normally agar is used as a solidifying agent to solidify the media. Another gelling agent named ‘Ficoll’ showed a good response in anther culture. The purity status of ‘Ficoll’ is much better compared to agar (Kao 1981). Ficoll is the synthetic polymer of sucrose and it has non-ionic status. It enhances the ratio of green plants to albino plant achievement (Lashermes 1992).

Rice anther should be collected from the middle to lower part of the donor plant (Jacquard et al. 2006). For pollen development, optimum nitrogen content should be main-tained. Optimum nitrogen content ensures quality micro-spores with high embryogenic potential (Lapitan and Violeta 1999). Temperature variation in the booting stage adversely affects the microspore development process (Lapitan and Violeta 1999; Mishra and Rao 2016). The role of anther orientation had been found critical in rice anther culture (Yang and Zhou 1979). Even though, anther from rationed rice had utilized in anther culture to assess the efficacy (Fig. 2, Guzmán and Arias 2000).

A great extent of work has been done in rice haploid and double haploid research. China is holding a strong position having more than 100 DH lines or varieties developed through anther culture. Likewise, other few countries like India, Japan, South Korea, Hungary have devolved DH lines using the anther culture (Siddique 2015).

As mentioned before, anther and microspore cultured haploid and DH has used to enhance the basic research, quantitative, qualitative, resistance status, and so on in rice. The first salt-tolerant indica rice variety named PSBRc50 ‘Bicol’ was developed by anther culture and released for cultivating in the salinity-prone area (Senadhira et al. 2002). Then, in 2003 salinity tolerant DH rice variety released utilizing six F1 hybrids where indica and japonica varieties were exploited (Lee et al. 2003). A DH line named AC1 found from crosses of multiple salt-tolerant lines showed significant salinity tolerance. AC1 was found suitable for cultivating in saline-affected regions of Bangladesh (Thomson et al. 2010). Afterward, drought is another menace for rice production. A rice variety named “Janka” developed through anther culture having drought tolerance in India (Pauk et al. 2009). Likewise, cold tolerance at an early stage found in an Indian variety known as “Abel” has developed through anther culture (Pauk et al. 2009). Also, anther culture has successfully derived indica rice variety tolerant to aluminum toxicity (Dewi et al. 2009). Likewise, the DH rice line has developed having multiple stress tolerance; tolerant to blast, aluminum toxicity, and shade (Purwoko et al. 2010).

Hybrid rice could outyield commonly cultivated rice varieties (https://www.irri.org/hybrid-rice). But the yield potential of the hybrids depends on the purity of the parent line. But with the elapse of time, the parental line deter-iorates due to various factors and ultimately results in lower quality and yield (Mishra and Rao 2016). A restorer line of a hybrid named Minghui 63 purified through anther culture and showed significant results in yield and resistance (Bai et al. 1991; Zhu et al. 1998). Anther culture utilized to develop restorer (B) lines for cytoplasmic male sterile (A) line (Wang et al. 1994). Then, Palanisamy et al. (2019) developed an improved R line with superior grain quality.

Then, a linkage map has developed in rice using the DH line. These maps were utilized to identify the molecular markers linked with the important genes which had significant resistance against rice blast, bacterial blight, and sheath blight disease (Wang et al. 2001). Moreover, the DH line assisted to develop a chromosome map of rice (Forster and Thomas 2005). Significant quantitative cha-racters of rice are controlled by quantitative trait loci (QTL) (Datta 2005). The individual effect of QTL is very trivial and it is affected greatly by environmental factors. To overcome the issue, multiple replicated trials are needed to achieve the actual phenotype. As DH has true-breeding nature and convenience of producing large numbers, the DH plant is widely used for the identification and mapping of QTLs in rice (Mishra and Rao 2016). Sheath blight resistance-related QTL has been identified in japonica rice by using the DH population (Xu et al. 2011). Afterward, six brown planthoppers resistance-related QTL successfully mapped using the DH mapping population (Soundararajan et al. 2004). Likewise, 5 panicle layer-uniformity related, 2 yields, and yield component related QTLs were identified through exploiting the DH population (Ma et al. 2009). Additionally, rice root traits QTLs, plant growth, yield, and yield components QTLs were also mapped by utilizing the rice DH population (Li et al. 2003; Hittalmani et al. 2002).

Amylose content, protein content, and lipid content are the major rice quality indicators. Four amylose content related, 2 protein content-related, and 2 lipid content related QTLs had mapped using the DH rice population (Lee et al. 2014). Alongside, Anther culture has a great potentiality in speed breeding and grain quality enrichment (Xa and Lang 2011). A rice variety named Koshihikari gave superior grain and eating quality developed through DH technology (Xa and Lang 2011). Likewise, Milyang 90, Guan 18 are another culture-derived genotype that showed superior grain quality. Notably, Guan 18 is a hybrid indica rice variety (Chung 1987; Zhu and Pan 1990).

In mutation breeding, Gamma irradiated rice anther lines gave an early and higher number of flower-producing plants (Myint et al. 2005). Then, mutant double haploid plant resistance to Striga (a parasitic weed of rice) was achieved through anther culture (Table 1) (Shariatpanahi et al. 2018).

Iron and Zinc deficiency is a common problem around the world that leads to various avoidable diseases. Bio- fortification of the major cereals like rice is one of the probable solutions. Plant breeders are trying to develop new indica rice varieties enriched with zinc and iron by utilizing japonica donors. International Rice Research Institute had started a project to attain the goal and DH production through anther culture had considered as one of the probable approaches. IRRI had evaluated above 1500 DH lines to find the potential lines enriched with micro-nutrients like iron and zinc till 2009 (Grewal 2009; Mishra and Rao 2016).

Then, the DH population have utilized in marker-assisted selection (MAS) breeding considering the phenotypic precision (William et al. 2002). A maintainer line has developed through the coupled utilization of anther culture and MAS. Notably, the cytoplasmic sterile maintainer had disease resistance, aroma, and red hull. Then, using this same approach dwarfing gene and glast resistance gene had introgressed in Basmati rice cultivar named ‘Ranbir Basmati’ (Samal et al. 2019). Afterward, anther culture- based Agrobacterium-mediated genetic transformation had conducted in rice (Chen et al. 2006; Li et al. 2007, 2010).

Wheat perspectives

In 1973, the first anther culture-derived bread wheat (T. aestivum) plant regenerated by three research groups (Ouyang et al. 1973; Chu et al. 1973; Picard and De Buyser 1973). Likewise rice, spontaneous doubling prevails in bread wheat ranging from 25 to 70% (Maluszynski et al. 2003a, b).

The first DH wheat variety was developed by China in 1985 named ‘Jinghua No-1’ (Hu et al. 1985). Within a short time, a DH variety named Florin developed in France (De Buyser et al. 1987). In anther culture-driven DH wheat cultivar development, China showed good progress. Accord-ing to Hu (Hu 1997), china developed 21 wheat cultivars until 1991 utilizing anther culture technique. Likewise, other countries developed DH wheat varieties and among those “Gk Delibab” (Pauk et al. 1995), ‘SV Agaton’ (Tuvesson et al. 2003), ‘McKenzi’ (Graf et al. 2003), or ‘AC Andrew’ (Sadasivaiah et al. 2004), ‘Huapei 8’ (Ming- hui et al. 2011), ‘Kharoba’ (Elhaddoury et al. 2012) and ‘GK Déva’ (Pauk et al. 2020) are good examples. In addition, Hume, Gregory, Gladius, Axe, Crusader, Spitfire, Cobra, Gauntlet, Merlin, Fang, and Espada are few DH wheat varieties (Broughton et al. 2014).

In Canada, 9 classes of Western Canadian Wheat are prevailing considering the milling quality (https://www.grainscanada.gc.ca/en/grain-quality/grain-grading/wheat-classes.html). Among those classes, Canada Western Red Spring (CWRS) is one. According to Dunwell (2010), 3 out of 5 wheat cultivars of CWRS were DH variety and a cultivar named “Lillian” (double haploid) covered 15% area of CWRS. Then “AC Andrew” named wheat cultivar covered the 99% cultivating area of Canada Western Soft White Spring (CWSWS) class which was also DH variety (Dunwell 2010).

Factors affecting the wheat anther culture had thoroughly stated in Lantos and Pauk (2020). In brief, wheat has a strong genotypic dependency for in vitro androgenesis. Re-searchers had identified numerous wheat varieties amenable to anther culture (viz ‘Chris’, ‘Pavon’, ‘Svilena’, ‘Bob’) which become the desired test genotype (Lazar et al. 1984; Lantos et al. 2013; Castillo et al. 2015; Nielsen et al. 2015; Seifert et al. 2016). In addition, few recalcitrant varieties have also been identified (Torp et al. 2001; Lantos et al. 2013; Castillo et al. 2015). According to Lantos and Pauk (2020), both the amenable and recalcitrant genotypes will assist in developing a new protocol to mitigating the undesirable effect of genotype dependency and genotype × treatment interactions. Generally, two types of bread wheat are cultivated globally depending on the environment; winter wheat and spring wheat. Spring wheat shows more responsiveness compared to winter wheat during anther culture (Weigt et al. 2020).

The growing condition of the donor plant is another deciding factor. Donor plants can be grown in controlled condition like a greenhouse year-round (Ghaemi et al. 1995; Torp et al. 2001; Pauk et al. 2003; Tuvesson et al. 2000, 2003; Soriano et al. 2007, 2008; Broughton 2008, 2011; Redha and Suleman 2011; Brew-Appiah et al. 2013; Sanchez-Diaz et al. 2013). Alternatively, donor plants can be raised in field condition. Considering efficacy, a field growing plant is ideal for large scale production where controlled condition grown plants are helping researchers to optimize the protocol and excel applied research year-round (Pauk et al. 2003; Chauhan and Khurana 2011; Lantos et al. 2013; Zhao et al. 2015, 2017; Weigt et al. 2016, 2019; Lazaridou et al. 2017; Lantos and Pauk 2020). Mid- to late uninucleate stages of the microscope are the suggested stage for culturing anther (Soriano et al. 2007, 2008; Broughton 2008, 2011; Chauhan and Khurana 2011; Redha and Suleman 2011; Rubtsova et al. 2013; Sanchez- Diaz et al. 2013; Zhao et al. 2015; Castillo et al. 2015; Echávarri and Cistué 2016; Weigt et al. 2016, 2019; Lazaridou et al. 2017; Broughton et al. 2020; Orlowska et al. 2020). Alternatively, early- and mid-uninucleate micro-spores were also utilized for wheat anther culture (Datta and Wenzel 1987; Datta and Wenzel 1987, 1995; Tuvesson et al. 2000, 2003; Datta 2005; Lantos et al. 2013; Lantos and Pauk 2016). Various stress factors like cold, heat, starva-tion, colchicine, osmotic shock, 2-HNA, DMSO, etc. had found beneficial (Liu et al. 2001; Barnabás 2003; Shariatpanahi et al. 2006; Echávarri and Cistué 2016). Short time cold treatment (3-8 days, 4-6℃) (Ghaemi et al. 1995; Broughton 2008, 2011; Rubtsova et al. 2013; Zhao et al. 2015, 2017; Weigt et al. 2016, 2019; Lazaridou et al. 2017; Sen 2017), long-duration cold treatment (2-5℃, 10 days-4 weeks) (Pauk et al. 2003; Lantos et al. 2013; Lantos and Pauk 2016; Coelho et al. 2018; Wang et al. 2019), starvation (solely or combined with chemical treatment) (Soriano et al. 2007, 2008; Sanchez-Diaz et al. 2013; Castillo et al. 2015; Echávarri and Cistué 2016), heat treatment (3 days, 32℃) has implied successfully (Ouyang et al. 1983; Pauk et al. 2003; Shariatpanahi et al. 2006; Lantos et al. 2013; Lantos and Pauk 2016). Previous studies indicated that colchicine has the capability to prompt microspore division and also can excel in the gametic embryogenesis in wheat (Barnabas et al.1991). A similar effect has been found in the case of Brassica (Mollers et al. 1994), rice (Alemano and Guiderdoni 1994).

Various culture media has been used in wheat anther culture such as AM, C17, P2, P4, LIM, W14, MS3M (Lantos and Pauk 2020). Among those W14, and MS3M media have been using frequently (Ouyang et al. 1989; Lantos et al. 2013; Rubtsova et al. 2013; Lantos and Pauk 2016; Lazaridou et al. 2017; Zhao et al. 2017; Soriano 2007; Sanchez-Diaz et al. 2013; Castillo et al. 2015; Echávarri and Cistué 2016). Lantos and Pauk (2020) have claimed that the modified W14 medium has found effective in wheat anther culture which has strengthened by previous findings (Lantos et al. 2013, 2018, 2019; Lantos and Pauk 2016; Kanbar et al. 2020; Pauk et al. 2020). Then, as a carbon source maltose has used frequently. The ficoll has been utilized as an osmotic agent (Hunter 1987; Datta and Wenzel 1987). Various growth regulators (2,4-D, benzyl adenine, centrophenoxine, dicamba, indole-3-acetic acid, kinetin, etc.) also exploited successfully to induce wheat plantlet (Fig. 2; Lantos and Pauk 2020; Castillo et al. 2015; Weight et al. 2016, 2019; Zhao et al. 2015, 2017; Orlowska et al. 2020; Rubtsova et al. 2013). In addition, a supplement like exogenous aliphatic polyamines (putrescine, cada-verine, spermidine, and spermine) was found beneficial including several wheat cultivars (Mishra and Rao 2016; Rajyalakshmi et al. 1995).

Afterward, as plant regeneration media, 190-2, J25-8, and MS media proved their effectiveness (Lantos and Pauk 2020; Lantos and Pauk 2016; Lazaridou et al. 2017; Orlowska et al. 2020; Castillo et al. 2015; Echávarri and Cistué 2016; Zhao et al. 2015, 2017; Weigt et al. 2016, 2019). A modification of anther culture named “shed culture” had also practiced in wheat. Here anther is sti-mulated to early dehiscence and after that, the freed micro-spores are cultured in a liquid medium having high osmo-larity (Touraev et al. 1996b, 1997).

Utilizing the DH technique, QTL identification and genetic mapping of various traits of wheat have been done. Photoperiod, plant height, flowering time, yield compo-nents, and grain yield-related QTL has mapped using DH line (Sourdille et al. 2000; Heidari et al. 2012; Cuthbert et al. 2008; Kuchel et al. 2007a, b). Alongside, Fusarium Head Blight (FHB), Septoria tritici blotch resistance QTL had identified and mapped through DH (Suzuki et al. 2012; Kelm et al. 2012). Then, 6 lodgings resistance-related QTL has unveiled by utilizing DH mapping population of wheat developed by anther culture (Hai et al. 2005). Then, QTL related to nitrogen uptake capacity also unfolded by DH mapping population of wheat developed by anther culture (An et al. 2006). Then, 6 salt tolerance QTL had found where the mapping population was a cross combination of Excalibur and Kukri wheat variety (Table 2) (Asif et al. 2018).

Additionally, the DH wheat plant has the potentiality to develop hybrid wheat varieties (Longin et al. 2014). A restorer line of hybrid wheat was bred using anther culture (Shi-kuan and Ya-ying 1985). Similarly, Shimada et al. (1994) developed a male sterility-maintainer line of wheat hybrid. In the case of genetic transformation, few reports had found where anther culture-derived haploid wheat embryo or callus had been used as an explant (Haliloglu et al. 2004; Chauhan and Khurana. 2011; Rustgi et al. 2020). Likewise, target mutagenesis was successfully achieved by genome editing utilizing the wheat microspore. The sgRNA of CRISPR/Cas9 had been delivered to wheat microspore and calli by electroporation and Agrobacterium-mediated transformation (Bhowmik 2018; Liu et al. 2020). Even though, for the first time in genome editing, the direct delivery of the Zinc finger nuclease (ZFN) protein in the intact wheat microspore had been claimed by Bilichak et al. (2020). They used purified ZFN along with the cell‐penetrating peptides (CPP) to deliver to the microspore where the microspore wall remains intact.

Brassica perspectives

Brassica is a large genus having species including oil-seed and vegetable crops (Ferrie et al. 1999; Barro and Martin 1999; Chuong et al. 1988). First Brassica haploid was achieved in B. oleracea (Kameya and Hinata 1970). Afterward, Keller et al. (1975) and Thomas and Wenzel (1975) succeeded in haploid regeneration in B. campestris and B. napus respectively. In 1982, haploid B. napus achi-eved from microspore culture (Lichter 1982). Notably, in B. napus, it has been estimated that microspore culture is tenfold efficient than the anther culture (Siebel and Pauls 1989). Especially, double haploid production through microspore culture has effectively used in B. rapa, B. oleracea, B. napus, B. carinata, and B. juncea and numerous varieties had developed in B. napus and B. oleracea (Watts et al. 2020) and numerically it is above 50 (Ferrie and Möllers 2011).

Various factors have a direct effect on haploid and double haploid production of Brassica and the principal factors had stated briefly by Watts et al. (2020). Especially for white cabbage, the responsible factor had been opti-mized for the ease of microspore embryogenesis and double haploid production (Bhatia et al. 2021). Firstly, Brassica has a genotypic dependency. Notably, the geno-type dependency could be transferred from a highly respon-sive to a low responsive genotype. Especially, the respon-sible loci for embryogenesis have been identified in B. napus and B. campestris (Cloutier et al. 1995; Ajisaka et al. 1999). Then, the growth environment of the donor plant has a great role in anther or microspores derive embryogenesis (Watts et al. 2020). In most cases, plants are grown at low temperatures responded significantly in Brassica micro-scope embryogenesis (Watts et al. 2020).

The early uninucleate to early binucleate pollen gave the best response (Gil-Humanes and Barro 2009). Individually, B. napus are good at the late uninucleate stage (Kott et al. 1988). Bud size is also a deciding factor for achieving the best response and species-wise variation is common. Optimum size should be identified for getting the best response (Gil-Humanes and Barro 2009; Watts et al. 2020). Then, Pretreatment with various stressors has found beneficial. Both cold and heat treatment had implied in Brassica microspore culture (Watts et al. 2020; Murovec and Bohanec 2011). Especially, for Brassica heat treatment is considered as an essential factor (Murovec and Bohanec 2011). Around 32℃ ranging from 1 to 4 days is mandatory for embryo regeneration of B. napus (Pechan and Keller 1988), B. carinata (Chuong and Beversdorf 1985), and B. juncea (Ohkawa et al. 1988), and B. oleracea (Takahata and Keller 1991). Noteworthy, heat stress has an anta-gonistic effect on microtubule development (Watts et al. 2020).

The modified Nitsch basal medium (Lichter 1982) and B5 medium are commonly used in Brassica (Gil-Humanes and Barro 2009). Sucrose is commonly used as the osmoticum and carbon source, as well as l-glutamine and l-serine, which serve as organic nitrogen sources (Licher 1982; Baillie et al. 1992; Babbar et al. 2004). Growth hormones had found beneficial in few genotypes, for instance, the application of abscisic acid increases embryo achievement in B. oleracea (Rudolf et al. 1999). Addi-tionally in B. rapa, B. oleracea, and B. napus (Guo and Pulli 1996; da Silva Dias 1999; Kott and Beversdorf 1990) activated charcoal was used and a beneficial response has perceived (Watts et al. 2020).

As the rate of spontaneous double haploid induction is very low in Brassica, various chromosome doubling agents, for instance, colchicine, oryzalin, amiprophosmethyl, tri-fluralin, and propanamide are utilized to achieve a double haploid plant. Amongst, colchicine is widely utilized and factors like concentration, duration of treatment, and plant developmental stage are considered (Watts et al. 2020). Notably, around 50-500 mg/L colchicine is used for 15-24 hours duration to yield double haploid (Watts et al. 2020; Mollers 1994; Zhou et al. 2002).

The double haploid Brassica lines have also been utilized to assess breeding potential, develop mapping population, QTL analysis, linkage mapping, and whole-genome se-quencing (Table 2; Sing et al. 2021; Wang et al. 2011; Yang et al. 2016; Pink et al. 2008). Especially in mutation breeding, microspore has been used as the inception medi-um of mutation. After mutation induction, the changed segment of DNA, even if a recessive trait will be altered into a homozygous state due to the double haplodization (Watts et al. 2020). In Brassica, several reports have found where microspore-based mutations were conducted. A tho-rough discussion regarding microspore embryogenesis had discussed by Ferrie and Möllers (2011). Notably, Ferrie et al. (2008) treated the microspore of B. rapa, B. napus, and B. juncea with a mutagenic agent and developed Brassica lines with altered fatty acid content. Likewise, high oleic acid-containing Brassica was also achieved (Turner and Facciotti 1990; Wong and Swanson 1991). Then, a similar approach was used to locate lower glucosinolate containing B. napus germplasm (Burbulis et al. 2001). Afterward, cold tolerant, salt-tolerant, and sclerotia-resistant mutant lines have developed through this In vitro mutagenesis approach (Rahman et al. 1995; Liu et al. 2005; McClinchey and Kott 2008).

In genetic transformation; Agrobacterium tumefaciens- mediated transformation (Pechan 1989; Dormann et al. 2001; Cegielska-Taras et al. 2008), particle gun bombard-ment (Fukuoka et al. 1998; Nehlin et al. 2000), micro-injection (Jones-Villeneuve et al. 1995), and electroporation (Guerche et al. 1987; Jardinaud et al. 1993) microspore utilized as the recipient of the transgene. Also, coupled utilization of biolistic delivery along with secondary em-bryogenesis was also reported in the microspore-based transformation of Brassica (Table 3) (Abdollahi et al. 2009, 2010).

Tobacco perspectives

Tobacco is considered one of the Rosetta plants studied extensively in plant biology (Belogradova et al. 2009). After Datura innoxia, tobacco (Nicotiana tabacum) is the second species where anther culture had implied success-fully to achieve haploid and double haploid plant (Guha and Maheshwari 1964; Bourgin and Nitsch 1967; Nitsch 1968; Nitsch and Nitsch 1969; Nitsch 1971, 1972).

Tobaccos responsiveness to anther culture is compara-tively very high and demands less complexity (Sunderland 2012). Several factors like donor plant growth condition, genotype, age, pre-treatment, photoperiod difference, anther developmental stage, and culture condition have a direct influence in increasing embryogenic response in tobacco (Dunwell 1976; Belogradova et al. 2009). Then, the pre-treatments such as chilling of buds, anaerobic anther treat-ment, and pre-treatment of the anther in water-saturated condition had found effective (Nitsch and Norreel 1973; Duncan and Heberle 1976; Sunderland 1978; Imamura and Harada 1981; Dunwell 1981). As culture media, N6 (Chu, 1978), MS (Murashige and Skoog 1962), H (Nitsch 1972) media were utilized and Sunderland (2012) recommended the N6 media. Especially, the presence of iron in culture media is vital because lacking iron inhibits embryogenesis in the globular stage (Nitsch 1972). Using activated charcoal in the medium found beneficial in previous studies. The charcoal absorbs the inhibitory and toxic substances from medium or agar or from the wall of the senescing cell (Anagnostakis 1974; Kohlenbach and Wernicke. 1978; Horner et al. 1977).

Generally from one anther of tobacco 1-135 plantlet could be achieved (Sunderland and Wicks 1971; Nitsch 1972; Horner et al. 1977). The growth stage of pollen is important. Empirically, it has been observed that bi-cellular pollen just after completing its first mitotic division gave consistent results (Sunderland and Wicks 1969; Sunderland 2012). Another, corolla length of tobacco could give an inkling about the anther responsiveness and the favorable range between 15 and 25 mm (Suntherland 2012).

Around 10-12% of plants derived from tobacco micro-spore culture are spontaneous DH. Then, colchicine could be to the diplodizing tobacco plant (Touraev and Heberle- Bors 2003). The ploidy level can be identified using flow cytometry or chromosome counting by cytological proce-dure (Touraev and Heberle-Bors 2003). Overall, the detailed procedure of tobacco anther and microspore culture has been stated thoroughly in Touraev and Heberle-Bors (2003) and Suntherland (2012). Especially, Sunderland (2012) stated the protocols of developing plantlets from tobacco anther culture. The first protocol Involves using a solid agar medium and the second one utilizes liquid media where float culture is practiced and pretreatment is man-datory (Sunderland 2012).

Anther culture had implied in tobacco to derive blue mold (Peronospora tabacina) black shank (Phytophthora parasitica var ‘nicotianae’) and tobacco mosaic virus- resistant variety and line (Nichols and Rufty 1992; Garcia et al. 1999). Likewise, Alternaria alterata, Powdery Mildew (Erysiphe cichoracearum) resistant line had found by utilizing the anther culture (Zhongxin et al. 1994; Gencer 2002). Mutant tobacco plants had developed through irra-diating the anther (Tong and Jia 1991; Daoru and Xinghua 1991). Genetics study of tobacco also carried out utilizing anther culture-derived tobacco plant (Fig. 2, Zhenji 1981; Ai and Chen 1981; Cho and Chang 1984). The chromoso-mal behavior was studied from the tobacco diploid ach-ieved by diploidization of the anther culture-derived haploids (Paz et al. 1994).

In genetic transformation, maize cytokinin-specific b- glucosidase gene was transferred to tobacco to observe the activity of b-glucosidase. Results showed that the overac-tivity of b-glucosidase dwindled the anther culture-derived pollen embryo regeneration (Dubová 1996). Similarly, using CRISPR/Cas9 based genome editing, a genetically fixed double haploid line had achieved. The microspore of tobacco and the leaf explant was the recipient of the sgRNA specifically constructed for the green fluorescent protein (GFP) gene (Table 4) (Schedel et al. 2017).

Albinism and the possible remedies

Albinism is one of the obstacles which act as a barrier in anther and microspore culture of plants. Albinism is a phenomenon where the regenerated plant could not run the photosynthesis process because of the lacking of chloro-plast and it hinders chlorophyll and other pigments like phytoene production (Makowska and Zimny 2015; Dunford and Walden 1991; Kumari et al. 2009; Liu et al. 2007; Qin et al. 2007). Albinism is inherently linked with the anther culture and microspore culture (Makowska and Zimny 2015). Review articles of Kumari et al. (2009) and Torp and Andersen (2009) tried to state important issues regard-ing albinism in plants.

Though there is no deep and clear indication about albinism occurrence, many researchers tried to unfold the behind mystery (Makowska and Zimny 2015). Scientists attempted to explain it by cytological study, plastid genome study, and nuclear genome study (Makowska and Oleszczuk 2014). Especially from the cytological aspect, the struc-tural change of chloroplast at various stages of anther or microspore culture which resist the chloroplast genesis by comparing the in vitro derived green plant and albino plant (Caredda et al. 1999, 2000, 2004). Likewise, it has attempted to find out the difference in the plastid genome and nuclear genome by comparing the green and the albino plant (Hofinger et al. 2000; Muñoz-Amatriaín et al. 2008, 2009; Makowska and Zimny 2015). A thorough discussion re-garding this research area had done by Makowska and Oleszczuk (2014). Apart from these assumptions, Duarte- Aké et al. (2016) tried to describe albinism from the epigenetics view. In brief, they found three types of plantlets (green, variegated, and albino) by culturing the somaclones of Agave angustifolia Haw and observed different types of DNA methylation patterns. Finally, they concluded that in vitro culture induces “Epigenetic Stress Memory” which might favor the development of the albino and variegated shoot by a chromatic shift.

Genotypic variation has a direct role in albino plant occurrence. For example, in winter barley the frequency of albino plant occurrence is lower than the spring wheat. This phenomenon could be explained by the evolutionary adap-tation as the winter one gone through low-temperature stress (Caredda et al. 2000; Castillo et al. 2000; Makowska and Zimny 2015).

To overcome albinism many attempts had taken and those empirical studies suggested the calibration of the parameters in in vitro anther culture. Among those, donor plant culturing environment, the growth stage of micro-spore, pretreatment, culture method, and media composi-tion is notable (Makowska and Zimny 2015). Similarly, Sriskandarajah et al. (2015) stated that optimization of the culture method will help to reduce this phenomenon, as it has a genotypic dependency.

For instance, cytokinins like thidiazuron and dicamba are used in induction media, and meta-topoline is utilized in regeneration media in barley microspore culture. The result showed that albino plant achievement dwindled signifi-cantly (Esteves et al. 2014). Afterward, Calic et al. (2012), showed that abscisic acid has a significant role in achieving green plant over the albino plant in horse chestnut (Aesculus hyppocastanum). Then, Mannitol along with colchicine or DMSO gave a higher number of green plantlets and DH plants compared to the starvation solely induced by mannitol (Soriano et al. 2007; Echávarri and Cistué 2016).

As mentioned, there is no clear and in-depth under-standing of the albino plant formation in androgenesis. Makowska and Oleszczuk (2014) stated the plastid behav-ior during androgenesis. They inferred that due to the failure of the reprogramming from gametophyte to sporo-phyte, chloroplast might not develop. Likewise, they inferred that early-stage microspore might reduce the albino plant but no findings ossified this assumption. Then, still, it is unknown when the plastid genome was modified that leads to albino plants (Makowska and Oleszczuk 2014). A few numbers of QTL had identified in the nuclear genome which has a direct effect on the green and albino plant ratio. Still now this number is very trivial (Yamagishi et al. 1998; Makowska and Oleszczuk 2014).

So, it is evident that unfolding the mechanism of albino plant formation is the important part. This could forward the way of androgenesis research. Notably, researchers are trying to explain it from the epigenetic aspect. In the future, the extensive epigenetic study will give pace to fathom albinism.

CONCLUSION

Androgenesis is successfully exploited for more than 5 decades and many haploid and doubled haploid plants had regenerated for basic and applied research. Most impor-tantly, various crop cultivars (barley, wheat, rice, Brassica, etc) have been cultivating around the world which has de-veloped through androgenesis. By 2050, the world popula-tion will be around 9.7 billion (https://www.worldometersinfo/world-population/). Additionally, the intensity of bio-tic and abiotic stresses will threaten the global food produc-tion. To tackle these issues, a more updated bio-fortified, climate-resilient, disease, and pest-resistant cultivar is needed. Double haploid developed through androgenesis will be one of the important tools to achieve it. The haploid and double haploid will be con-tinued to contribute to gene mapping, QTL mapping, Genome-Wide Association Study (GWAS), mutation breed-ing, Marker Assisted Selection (MAS), Marker-Assisted Backcross (MAB) breeding. Similarly, in genetic transfor-mation and genome editing, it will be exploited, especially in the lines where the zygotic embryo is recalcitrant to receive the trans-gene or sgRNA. Finally, to dissect the phenomenon of albinism, the scien-tist will try to utilize epigenetics extensively along with the existing approaches. Finally, cutting-edge techniques like the reverse genetic approach, genome editing especially CRISPR/Cas9 will be a game-changer.

Fig. 1

Schematic diagram showing the anther culture technique.

Fig. 2

Embryogenic calli (a), Embryo Like Structure (ELS), (b) and plant regenerating (c) from ELS of rice anther culture (Guzmán and Arias 2000); ELS (d, e), regenerating plantlets (f), regenerated plantlet (g), haploid (n) and double haploid (2n) wheat plant (h) by anther culture (Rubtsova et al. 2013); green and albino plantlet achieved through barley anther culture (i) (Sriskandarajah et al. 2015); Segregating green and albino plantlet found by tobacco anther culture (j) (Dunwell 2010).

Table 1

List of rice varieties developed through anther culture having stress tolerance and other agronomic special features (Adopted from Sarao and Gosal et al. 2018).

Table 1
Variety name	Characteristics	Country	Reference
Huayu I, Huayu II, Xin Xiu, Late Keng 959, Tunghua 1, Tunghua 2, Tunghua 3, Zhonghua 8, Zhonghua 9, Huahanzao, Huajian 7902, Tanghuo 2, Shanhua 7706, Huahanzao 77001, Nanhua 5, Noll, Hua 03	Rice blast and bacterial blight resistance along with quality grain	China	Zang 1980; Hu and Zeng 1984; Chen 1986; Loo and Xu 1986; Yang and Fu 1989
Guan 18	Early maturity; good quality and disease resistance	China	, Zhu and Pan 1990
Huayu 15	Resistant to lodging and diseases; good quality	China	, Shouyi and Shouyin 1991
Milyang 90	Brown planthopper and stripe rust-resistant with better grain quality	China	, China Chung 1987
Hwacheongbyeo, Joryeongbyeo, Hwajinbyeo	Resistant to brown planthopper, rice stripe tenui virus, blast, and bacterial blight	China	, Lee et al. 1989
Risabell	High milling and cooking quality; resistant to blast	India	, Pauk et al. 2009
CR Dhan 10 (CRAC2221-43), Satyakrishna	Resistant to neck blast, sheath-rot, and yellow stem borer	India	CRRI Annual Report 2007–2008; Rao and Gosal 2018
CR Dhan 801 (CRAC2224-1041, IET18720), Phalguni	Resistant to leaf blast, gall midge; moderately resistant to sheath rot, rice stripe tenuivirus, yellow stem borer, brown spot, and sheath blight	China	CRRI Annual Report 2009–2010; Rao and Gosal 2018

Table 2

Double haploid technology in wheat breeding.

Table 2
Application	Trait	Reference
Development of cultivars	Improving flour quality, higher molecular weight glutenin, rust resistance	Ushiyama T 2008; Touraev et al. 2009; Bakhshi et al. 2012
Mapping quantitative trait loci	Foliar disease of wheat, important agronomic traits, Fusarium head blight, number of tillers, number of spikes per plant and spike length, leaf rust resistance, photoperiod response, heading time, crossability among different wheat genome	, Chu et al. 2008a, b; Yang et al. 2005; Li et al. 2010; Suenaga et al. 2005; Huang et al. 2003; Sourdille et al. 2000; Tixier et al. 1998
Production of transgenic plants	Salt tolerance, drought tolerance	, Chauhan and Khurana 2010, 2011; Khurana et al. 2011

Table 3

List of Brassica species with mapping traits using double haploid technology.

Table 3
Species	Trait	Reference
B. juncea	Seed color, Yield contributing QTL, Oil quality and oil content, White rust resistance, Seed glucosinolate, Seed weight	, Padmaja et al. 2014; Ramchiary et al. 2007a; Yadava et al. 2012; Jagannath et al. 2011; Panjabi-Massand et al. 2010; Rout et al. 2018; Ramchiary et al. 2007b; Dhaka et al. 2017b
B. oleracea	Seed vigour and pre-emergence seedling growth traits	, Bettey et al. 2000
B. napus	Seed glucosinolate quantity	, Uzunova et al. 1995

Table 4

Haploid and double haploid technology in tobacco breeding.

Table 4
Technology	Application	Reference
Haploid	Genome mapping, identification of quantitative trait loci (QTL), genetic transformation, marker assisted selection in breeding for new cultivars	, Wernsman 1993; Tai 2005; Milla et al. 2005
Haploid & double haploid	Environment-friendly F₁ hybrid system	, Ribarits et al. 2007

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Anther Culture in Crop Plants: Progress and Perspectives

Plant Breed. Biotech.. 2023;11(2):69-96. Published online June 1, 2023

DOI: https://doi.org/10.9787/PBB.2023.11.2.69

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Anther Culture in Crop Plants: Progress and Perspectives

Plant Breed. Biotech.. 2023;11(2):69-96. Published online June 1, 2023

DOI: https://doi.org/10.9787/PBB.2023.11.2.69

Figure

Anther Culture in Crop Plants: Progress and Perspectives

Fig. 1 Schematic diagram showing the anther culture technique.

Fig. 2 Embryogenic calli (a), Embryo Like Structure (ELS), (b) and plant regenerating (c) from ELS of rice anther culture (Guzmán and Arias 2000); ELS (d, e), regenerating plantlets (f), regenerated plantlet (g), haploid (n) and double haploid (2n) wheat plant (h) by anther culture (Rubtsova et al. 2013); green and albino plantlet achieved through barley anther culture (i) (Sriskandarajah et al. 2015); Segregating green and albino plantlet found by tobacco anther culture (j) (Dunwell 2010).

Fig. 1

Fig. 2

Anther Culture in Crop Plants: Progress and Perspectives

List of rice varieties developed through anther culture having stress tolerance and other agronomic special features (Adopted from Sarao and Gosal et al. 2018).

Variety name	Characteristics	Country	Reference
Huayu I, Huayu II, Xin Xiu, Late Keng 959, Tunghua 1, Tunghua 2, Tunghua 3, Zhonghua 8, Zhonghua 9, Huahanzao, Huajian 7902, Tanghuo 2, Shanhua 7706, Huahanzao 77001, Nanhua 5, Noll, Hua 03	Rice blast and bacterial blight resistance along with quality grain	China	Zang 1980; Hu and Zeng 1984; Chen 1986; Loo and Xu 1986; Yang and Fu 1989
Guan 18	Early maturity; good quality and disease resistance	China	Zhu and Pan 1990
Huayu 15	Resistant to lodging and diseases; good quality	China	Shouyi and Shouyin 1991
Milyang 90	Brown planthopper and stripe rust-resistant with better grain quality	China	China Chung 1987
Hwacheongbyeo, Joryeongbyeo, Hwajinbyeo	Resistant to brown planthopper, rice stripe tenui virus, blast, and bacterial blight	China	Lee et al. 1989
Risabell	High milling and cooking quality; resistant to blast	India	Pauk et al. 2009
CR Dhan 10 (CRAC2221-43), Satyakrishna	Resistant to neck blast, sheath-rot, and yellow stem borer	India	CRRI Annual Report 2007–2008; Rao and Gosal 2018
CR Dhan 801 (CRAC2224-1041, IET18720), Phalguni	Resistant to leaf blast, gall midge; moderately resistant to sheath rot, rice stripe tenuivirus, yellow stem borer, brown spot, and sheath blight	China	CRRI Annual Report 2009–2010; Rao and Gosal 2018

Double haploid technology in wheat breeding.

Application	Trait	Reference
Development of cultivars	Improving flour quality, higher molecular weight glutenin, rust resistance	Ushiyama T 2008; Touraev et al. 2009; Bakhshi et al. 2012
Mapping quantitative trait loci	Foliar disease of wheat, important agronomic traits, Fusarium head blight, number of tillers, number of spikes per plant and spike length, leaf rust resistance, photoperiod response, heading time, crossability among different wheat genome	Chu et al. 2008a, b; Yang et al. 2005; Li et al. 2010; Suenaga et al. 2005; Huang et al. 2003; Sourdille et al. 2000; Tixier et al. 1998
Production of transgenic plants	Salt tolerance, drought tolerance	Chauhan and Khurana 2010, 2011; Khurana et al. 2011

List of Brassica species with mapping traits using double haploid technology.

Species	Trait	Reference
B. juncea	Seed color, Yield contributing QTL, Oil quality and oil content, White rust resistance, Seed glucosinolate, Seed weight	Padmaja et al. 2014; Ramchiary et al. 2007a; Yadava et al. 2012; Jagannath et al. 2011; Panjabi-Massand et al. 2010; Rout et al. 2018; Ramchiary et al. 2007b; Dhaka et al. 2017b
B. oleracea	Seed vigour and pre-emergence seedling growth traits	Bettey et al. 2000
B. napus	Seed glucosinolate quantity	Uzunova et al. 1995

Haploid and double haploid technology in tobacco breeding.

Technology	Application	Reference
Haploid	Genome mapping, identification of quantitative trait loci (QTL), genetic transformation, marker assisted selection in breeding for new cultivars	Wernsman 1993; Tai 2005; Milla et al. 2005
Haploid & double haploid	Environment-friendly F₁ hybrid system	Ribarits et al. 2007

Table 1 List of rice varieties developed through anther culture having stress tolerance and other agronomic special features (Adopted from Sarao and Gosal et al. 2018).

Table 2 Double haploid technology in wheat breeding.

Table 3 List of Brassica species with mapping traits using double haploid technology.

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