1 INTRODUCTION
Maize (
Zea mays L.) is a crop of major importance in China. Approximately two-thirds of its annual output is used for livestock and poultry feed, with the remainder allocated to food uses and industrial processing, such as starch production (Huang et al.,
2026; Luo et al.,
2025). Therefore, improving maize yield and quality is crucial to safeguarding China's food security and supporting the growth of its livestock sector (Huang et al.,
2022). Anthocyanins belong to the flavonoid family and are water-soluble phenolic pigments. In nature, they typically exist as glycosides (Lu et al.,
2024). Anthocyanins display a diverse range of colors, which not only attract pollinators and enhance pollination efficiency but also scavenge oxygen free radicals in vivo, protect cells from oxidative damage, and thereby contribute to the prevention of cardiovascular diseases and the protection of human health (Krga and Milenkovic,
2019). In most modern maize varieties, anthocyanins accumulate only in a limited number of tissues, such as the basal leaf sheaths, or show virtually no accumulation (Petroni et al.,
2014). In high-anthocyanin maize (also referred to as purple, black, or red maize), the expression of anthocyanin biosynthesis-related genes is upregulated, resulting in substantial anthocyanin accumulation in multiple tissues, including kernels, husk leaves, stalks, tassels, and silks. The anthocyanin content in kernels typically ranges from 42 to 354 mg/100 g (Zhao et al.,
2008), and in some varieties, it can reach up to 1970 mg/100 g (Cai et al.,
2023), conferring high nutritional and health-promoting value.
Fresh-eating maize is characterized by its rich flavor and nutritional quality (Jiao et al.,
2025). In Chinese maize production, high-anthocyanin maize is frequently cultivated alongside other fresh-eating varieties, including sweet maize, waxy corn, sweet-waxy corn, and baby corn, increasing the economic returns of fresh-eating maize cultivation, enhancing its nutritional quality, and improving both its taste and health benefits (Li et al.,
2025a). Meanwhile, against the backdrop of rapid population aging in China, individuals aged 60 years and above accounted for more than 23% of the national population by the end of 2025 (National Bureau of Statistics). Developing functional varieties that enhance the intake of trace elements and functional nutrients among Chinese residents, and improve public health literacy, is therefore of positive significance (Li et al.,
2019a; Yan et al.,
2023). High-anthocyanin maize, with its rich nutritional profile and beneficial health-promoting properties, can help meet the population's growing demand for better health (Magni et al.,
2025).
As research on high-anthocyanin maize has progressed, the genetic regulatory network of anthocyanin biosynthesis has become increasingly clear. The theoretical foundation of high-anthocyanin maize has been substantially enriched, molecular breeding strategies have been continuously optimized, and breeding efficiency has significantly improved. This review summarizes the molecular genetic mechanisms of anthocyanins, anthocyanin extraction and purification, the health-promoting functions of high-anthocyanin maize, its genetic improvement, and its functional development and industrial application. It thus provides a theoretical basis for further elucidating the molecular regulatory mechanisms of high-anthocyanin maize, promoting the development of anthocyanin-rich maize food products, and advancing high-value industrial development.
2 Anthocyanin Biosynthesis and Molecular Genetics
2.1 Anthocyanin composition, coloration, and biosynthesis in maize
The basic structure of anthocyanins consists of two benzene rings (A ring and B ring) and a heterocyclic ring (C ring), with the C ring connected to the A ring and B ring via a three-carbon bridge. The diversity of anthocyanins is primarily determined by the number and position of glycosyl and hydroxyl groups at the R
1 and R
2 sites on the B ring, as well as by the aromatic acids and fatty acids linked to the glycosyl moieties (Fig. 1, Table 1). Anthocyanins, the glycosylated derivatives of anthocyanidins, constitute the predominant natural pigment form in plants and are directly responsible for the diverse coloration of plant organs. These structural variations give rise to six major types of anthocyanidins: cyanidin, pelargonidin, delphinidin, peonidin, petunidin, and malvidin (Xie et al.,
2024).
In maize kernels, anthocyanins are mainly localized in the vacuoles of the aleurone layer, scutellum, and pericarp tissues (Hu et al.,
2023). Both cyanidin and pelargonidin are present in the kernels of high-anthocyanin maize (Lu et al.,
2024). More than 22 anthocyanin glycosides have been identified in purple maize kernels, with cyanidin being the most abundant. In pink maize kernels, pelargonidin accounts for the highest proportion (Xie et al.,
2024). Delphinidin, petunidin, and malvidin are also present in maize, although at relatively low levels (Peniche-Pavía and Tiessen,
2020). The color of the yellow kernels most commonly found in commercial maize is determined by carotenoids, and these kernels contain only low levels of anthocyanins (Sánchez-Nuño et al.,
2024). Furthermore, in some maize varieties, the anthocyanin content in the husk leaves is up to ten times that in kernels, indicating that both kernels and other plant parts serve as important sources of anthocyanins (Cai et al.,
2023).
Anthocyanin coloration is modulated by pH, metal ion interactions, and structural modifications (Ye et al.,
2021). Because of their diverse hues, anthocyanins are widely utilized as natural pigments (Colombo et al.,
2021; Chatham et al.,
2019). In maize, anthocyanins exhibit pink coloration under acidic conditions, purple under neutral pH, and transition to yellow-green with concomitant rapid degradation under alkaline conditions (Barba et al.,
2022). Interactions of anthocyanins with metal ions such as iron and copper induce color changes, whereas antioxidants including vitamins C and E provide partial protection against oxidative damage (He et al.,
2026). Both ultraviolet radiation and elevated temperatures accelerate anthocyanin degradation, leading to discoloration of kernels and husks (Shi et al.,
2023; Zou et al.,
2023).
Anthocyanin biosynthesis in maize branches from the phenylpropanoid metabolic pathway (Liu et al.,
2021; Fang,
2015) and proceeds through five sequential stages: precursor synthesis, flavonoid backbone assembly, pigment molecular conversion, chemical modification, and vacuolar transport and localization (Lu et al.,
2023a). The biosynthetic process is described below in detail. First, phenylalanine is deaminated by phenylalanine ammonia-lyase (PAL) to yield cinnamic acid. Second, cinnamate 4-hydroxylase (C4H) introduces a hydroxyl group at the para position of the benzene ring, generating p-coumaric acid. Subsequently, 4-coumarate CoA ligase (4CL) activates p-coumaric acid through adenylation and thioesterification to produce p-coumaroyl-CoA (Mora et al.,
2022; Zhang et al.,
2025a). Next, chalcone synthase (CHS) catalyzes the condensation of p-coumaroyl-CoA with malonyl-CoA to form naringenin chalcone (Zhang et al.,
2018; Liu et al.,
2025), which is then converted to naringenin by chalcone isomerase (CHI). CHS and CHI are considered the key rate-limiting enzymes in the anthocyanin biosynthesis pathway (Jones et al.,
2017). Naringenin is further hydroxylated by flavanone 3-hydroxylase (F3H) to generate dihydrokaempferol, which is then catalyzed by flavonoid 3′-hydroxylase (F3′H) or flavonoid 3′,5′-hydroxylase (F3′5′H) to produce dihydroquercetin (the precursor of cyanidin) and dihydromyricetin (the precursor of delphinidin), respectively (Zhang et al.,
2018; Holly et al.,
2026). Dihydroflavonol 4-reductase (DFR) sequentially reduces dihydrokaempferol, dihydroquercetin, and dihydromyricetin to leucopelargonidin, leucocyanidin, and leucodelphinidin. These compounds serve as substrates and are converted, under the action of anthocyanidin synthase (ANS) followed by glycosylation catalyzed by UDP-glucose: flavonoid 3-O-glucosyltransferase (UFGT), into pelargonidin 3-glucoside, cyanidin-3-glucoside, and delphinidin-3-glucoside, respectively (Liu et al.,
2021; Kowalczyk et al.,
2024). These anthocyanin glucosides then undergo acylation and methylation catalyzed by acyltransferases, forming more stable anthocyanins. Finally, glutathione S-transferase (GST) mediates the conjugation of glutathione with cyanidin-3-glucoside (C3G) to produce glutathionylated C3G, which is transported into the vacuole via the glutathione conjugate transport system and a vesicle-mediated, non-carrier transport mechanism. The anthocyanins are ultimately stored as pigmented aggregates (Fig. 2) (Sunil and Shetty,
2022; Zhao et al.,
2024).
2.2 Regulatory genes of key enzymes in maize anthocyanin biosynthesis
In the maize anthocyanin biosynthetic pathway, multiple structural genes have been characterized through classical genetic mapping. Allelic variations directly determine the type, abundance, and tissue-specific distribution of pigment accumulation (He et al.,
2026).
C2 (
Colorless2) encodes chalcone synthase (CHS), which is expressed across multiple tissues and developmental stages (Han et al.,
2016). Structural disruption of the β-sheet motif in the
C2-encoded protein abolishes CHS activity, thereby preventing anthocyanin accumulation in kernels (Zhang et al.,
2018). As a gene encoding a rate-limiting enzyme,
C2 shows a significant positive correlation with anthocyanin content, and its downregulation markedly attenuates pigment deposition in transgenic plants (Nishihara et al.,
2005; Cai et al.,
2023).
Pr1 (
Purple aleurone1) encodes flavonoid 3′-hydroxylase (F3′H), which is a major determinant of anthocyanin chemical identity. In the homozygous recessive state,
Pr1 directs the accumulation of red pelargonidin-type anthocyanins, whereas the dominant homozygous allele enables 3′-hydroxylation, shifting synthesis toward purple cyanidin-type anthocyanins (Peniche-Pavía et al.,
2022). Thus, the allelic status of
Pr1 serves as a critical molecular marker for kernel color differentiation.
A1 (
Anthocyaninless1) encodes dihydroflavonol 4-reductase (DFR), an essential reductase acting downstream in the pathway. Introduction of a 35S promoter-driven
ZmA1 cDNA into anthocyanin-deficient mutants restores red pigment synthesis (Haselmair-Gosch et al.,
2018; Kumkum et al.,
2024).
A2 (
Anthocyaninless2) encodes anthocyanidin synthase (ANS), a 2-oxoglutarate-dependent dioxygenase that catalyzes the oxidation of colorless leucoanthocyanidins to colored anthocyanidins; mutations in
A2 block anthocyanin accumulation and alter plant pigmentation (Riaz et al.,
2019; Liu et al.,
2025).
Bz1 (
Bronze1) encodes UFGT (Li et al.,
2019b), and its expression is suppressed under high-temperature stress, leading to reduced pericarp anthocyanin accumulation (Li et al.,
2019b).
Bz2 (
Bronze2) encodes glutathione S-transferase (GST), which conjugates glutathione to anthocyanin precursors to facilitate vacuolar sequestration. Notably, cadmium exposure strongly induces
Bz2 expression, implicating this gene in metabolic reprogramming of anthocyanins under heavy metal stress (Table 2) (Marrs and Walbot,
1997; Li et al.,
2023).
2.3 Tissue-specific regulatory mechanisms of anthocyanin biosynthesis in maize
Tissue-specific distribution of anthocyanins in maize is not stochastic but is governed by distinct regulatory gene combinations.
C1 (
Colored aleurone1) predominantly functions in the kernel endosperm and aleurone layer. Heterologous expression of
ZmC1 in wheat triggers massive pigment production in coleoptiles, stems, and other vegetative organs (Riaz et al.,
2019), demonstrating its capacity to activate anthocyanin biosynthesis cross-species.
Pl1 (
Purple plant1), a paralog of
C1, encodes an MYB transcription factor. Despite high sequence similarity,
Pl1 exhibits a broader expression domain, regulating pigmentation in both floral and vegetative tissues while coordinately activating or repressing multiple key structural genes across flavonoid pathways (He et al.,
2026). EMS-based mutagenesis and CRISPR/Cas9 validation confirmed that
Pl1 is essential for anthocyanin biosynthesis in maize leaf sheaths, and overexpression of
Pl1 drives anthocyanin hyperaccumulation in leaf sheaths (Luo et al.,
2026).
R1 (
Colored1) and
B1 (
Colored plant1) are paralogous members of the bHLH transcription factor family and exhibit stringent tissue-specific expression patterns. Spatiotemporal expression analyses have demonstrated that the combinatorial interactions between the
C1/
R1 and
Pl1/
B1 gene pairs determine the spatial distribution of anthocyanins within maize kernels (Li et al.,
2019b). Specifically, the
Pl1/
B1 module governs anthocyanin accumulation in the pericarp, whereas the
R1/
C1 complex determines anthocyanin deposition in the aleurone layer, concomitantly elevating kernel anthocyanin contents to 1.2−1.8 mg/g. Furthermore,
Pr1 functions downstream of
C1 and
R1 to specifically modulate pelargonidin accumulation in the aleurone layer (Hichri et al.,
2011).
Allelic variation at
ZmR1 significantly influences anthocyanin accumulation across tissues. Cloning of
ZmR1 from inbred line CQ01 demonstrated that its overexpression confers intense purple pigmentation throughout the entire plant with substantially elevated anthocyanin levels (Luo et al.,
2022). This finding was applied in breeding practice through the development of the purple-kernel hybrid Jingke 968, derived from crossing the purple inbred line Jing 724 with Jing 92 (Luo et al.,
2022). Integrated transcriptomic and metabolomic analyses across kernel developmental stages reveal dynamic coupling among anthocyanin structural genes, regulatory transcription factors, and flavonoid metabolites, indicating that anthocyanin accumulation is governed by developmentally specific transcriptional programs rather than constitutive activation of the entire biosynthetic pathway (Table 3) (Ming et al.,
2021).
2.4 Assembly specificity and regulatory networks of the MBW complex in maize
Unlike the promiscuous assembly mode observed in Arabidopsis and other dicotyledonous species, where MBW complexes exhibit flexible combinatorial pairing (Gonzalez et al.,
2007; Goswami et al.,
2018), the maize MBW complex displays a highly simplified one-to-one assembly architecture. In this system, a single MYB (
C1 or
Pl1), a single bHLH (
R1 or
B1), and a single WD40 protein constitute the core regulatory unit (Paulsmeyer and Juvik,
2023). This streamlined configuration implies that maize anthocyanin regulation relies predominantly on the intrinsic spatiotemporal expression specificity of individual alleles rather than on the combinatorial diversity of complex components. Within this complex, the MYB transcription factor mediates specific DNA binding to target gene promoters, the bHLH transcription factor enhances transcriptional activity, and the WD40 protein functions as a scaffold to stabilize complex architecture (Lu et al.,
2023a; Luo et al.,
2022). The DNA-binding domain of maize
C1 recognizes chalcone synthase gene promoters (e.g.,
C2) via the GTR motif, exhibiting substantially higher binding affinity than that of conventional MYB proteins (Colombo et al.,
2021). WD40 proteins lack intrinsic DNA-binding capacity; instead, they indirectly modulate downstream gene transcription through mediating histone covalent modifications and participating in dynamic chromatin remodeling (Cai et al.,
2023).
PAC1 (
Pale aleurone color1) serves as the scaffold subunit of the maize MBW complex and directly activates structural genes to regulate anthocyanin biosynthesis (Yan et al.,
2019). However, the tissue-specific variation in
PAC1 expression levels and its quantitative contribution to complex stability remain systematically unexplored.
2.5 Molecular mechanisms of maize anthocyanin-environment interactions
Environmental effects on maize anthocyanin biosynthesis have been extensively documented, yet the molecular networks governing these interactions remain incompletely understood. Light is an important environmental cue that upregulates the expression of genes encoding key biosynthetic enzymes, including F3H and UFGT, leading to substantially enhanced anthocyanin accumulation in purple waxy maize kernels (Lu et al.,
2023b). Conversely, excessive light intensity induces DNA damage, thereby inhibiting anthocyanin synthesis (Guo et al.,
2008; He et al.,
2026). However, the precise mechanisms through which light signals activate the MBW complex via phytochrome and cryptochrome photoreceptors, as well as the molecular nodes connecting DNA damage response pathways to anthocyanin biosynthesis, remain to be defined. Temperature exerts significant effects on this metabolic pathway. Low-temperature conditions activate MBW complex-associated genes, resulting in marked upregulation of anthocyanin biosynthetic genes in maize seedlings. In contrast, exposure to 32°C significantly attenuates both anthocyanin synthesis and accumulation in kernels; elevated temperatures further enhance superoxide dismutase (SOD) activity, thereby accelerating anthocyanin degradation (Aguilar-Hernández et al.,
2019; Wang et al.,
2016). While downregulation of key structural genes such as UFGT under heat stress likely represents a direct cause of reduced pigment accumulation, the molecular mechanisms through which thermal signals are transduced to the MBW complex to suppress its transcriptional activation remain to be experimentally validated. Mineral nutrients additionally contribute to this regulatory network. Appropriate nitrogen application enhances the activity of anthocyanin biosynthetic enzymes and promotes pigment accumulation (Feng et al.,
2024). Increased soil zinc availability leads to vacuolar zinc enrichment, where Zn ions form stable complexes with anthocyanins, thereby enhancing pigment stability and promoting leaf anthocyanin accumulation (Utasee et al.,
2022; Janeeshma et al.,
2020). However, the specific binding sites, stability constants of these metallo-pigment complexes, and their consequent effects on vacuolar pH have yet to be precisely characterized in maize.
2.6 Unresolved issues in anthocyanin biosynthesis and molecular genetic research in maize
Although the basic framework of maize anthocyanin biosynthesis has been largely established, several critical gaps continue to constrain further advances in this field. First, the fine-scale regulatory logic underlying tissue-specific accumulation remains incompletely resolved. While anthocyanin deposition in the aleurone layer and pericarp is governed by the C1/R1 and Pl1/B1 regulatory modules, respectively, whether these two systems engage in cross-regulation remains unknown. Second, maize-specific mechanisms governing anthocyanin transport and vacuolar sequestration are poorly understood. In particular, the biophysical conditions driving pigment aggregate formation within vacuoles and their consequential effects on anthocyanin stability have received limited systematic investigation. Third, whether pyramiding high-anthocyanin alleles causes diversion of metabolic flux from primary metabolism into the phenylpropanoid pathway, thereby compromising grain filling efficiency, remains to be experimentally tested.
Leveraging the aforementioned molecular genetic regulatory networks, marker-assisted breeding enables the precise pyramiding of superior anthocyanin-associated alleles, facilitating the development of novel varieties with enhanced anthocyanin accumulation. Given maize's advantages of high productivity and wide geographic adaptability, increasing its anthocyanin content offers considerable potential for improving human health. The following section will detail the unique health benefits of high-anthocyanin maize from nutritional and biomedical standpoints.
3 Health-Promoting Effects of High-Anthocyanin Maize
3.1 Antioxidant effects
High-anthocyanin maize accumulates abundant anthocyanins in kernels and other tissues, with cyanidin- and pelargonidin-derived anthocyanins as the predominant components. Cyanidin-3-glucoside and related derivatives are important contributors to the antioxidant potential of purple maize (Cai et al.,
2023; Kim et al.,
2023). Their antioxidant activity is mainly associated with the polyphenolic structure of anthocyanins, which facilitates electron or hydrogen-atom donation and thereby contributes to free-radical scavenging (Hu and Xu,
2011; Sadowska-Bartosz and Bartosz,
2024).
Experimental studies using maize-derived anthocyanins support this antioxidant potential. Purified anthocyanins from corn have been reported to enhance antioxidant enzyme activity, reduce oxidative stress, and alleviate tissue injury in experimental models (Cui et al.,
2021; He et al.,
2024). These effects may be explained by three complementary mechanisms (Fig. 3): direct scavenging of reactive oxygen species (ROS), chelation of redox-active metal ions, and enhancement of endogenous antioxidant defenses. In particular, cyanidin-type anthocyanins can donate electrons or hydrogen atoms to reactive radicals and may also chelate metal ions through hydroxyl groups on the B ring. Mechanistic studies of cyanidin-3-glucoside and anthocyanin-rich preparations further suggest that Nrf2-related signaling may contribute to the activation of antioxidant and detoxification systems, thereby reducing oxidative stress and associated cellular damage (Xie et al.,
2018; Alharbi et al.,
2022; Ali et al.,
2024). Nevertheless, evidence from purified maize anthocyanins or isolated anthocyanin compounds should not be directly equated with the physiological effects of consuming whole high-anthocyanin maize. Anthocyanin stability, metabolism, food-matrix interactions, and bioavailability can influence their biological effects. Future studies should therefore link the anthocyanin profiles of specific maize genotypes with bioavailability and in vivo antioxidant outcomes to clarify how effectively high anthocyanin content translates into physiological benefits.
3.2 Anticancer effects
Acidified blue maize extracts exert antiproliferative effects against multiple tumor cell lines. Notably, cyanidin-3-(6''-malonylglucoside) I (CyMalGlu I) is a principal bioactive constituent, and its abundance negatively correlates with cancer cell viability (Urias-Lugo et al.,
2015). Anthocyanins have been shown to repair DNA damage-induced cellular carcinogenesis and promote cancer cell apoptosis (Stefanowicz-Hajduk et al.,
2021; Kowalczyk et al.,
2024). In studies on smooth muscle and liver cells, anthocyanins effectively inhibit lipid peroxidation and cytotoxic responses, reducing cellular DNA damage (Lazzé et al.,
2003). The anticancer effects of anthocyanins may involve the following four mechanisms. First, at the transcriptional level, they regulate the expression of key genes and proteins, arrest cell cycle progression, and induce cancer cell apoptosis (Ren et al.,
2003). Second, by inhibiting oxidative stress, they reduce DNA damage, prevent carcinogens from binding to DNA, and suppress the initiation of carcinogenesis (Singletary et al.,
2007). Third, they reduce the binding capacity of cancer cell receptors to the cell membrane, decrease the expression of cancer cell surface markers, and inhibit metastasis to other tissues and organs (Wu et al.,
2022). Finally, by reducing the energy supply required by cancer cells, they maintain the cells in a state of metabolic inhibition, ultimately leading to apoptosis (Herrera-Sotero et al.,
2019).
3.3 Cardiovascular protective and vision-protective effects
Purple maize anthocyanins, with cyanidin-3-glucoside (C3G) and its acylated derivatives as the principal bioactive constituents, exert cardioprotective effects through precise modulation of the AMPK/SIRT1/p53 signaling axis: they restore doxorubicin-induced aberrant AMPK activation, promote SIRT1-dependent p53 acetylation, and consequently downregulate pro-apoptotic effectors including cleaved-caspase 3, Puma, and p21, ultimately suppressing cardiomyocyte apoptosis (Cappellini et al.,
2025). Anthocyanins derived from other plant sources also exhibit similar bioactive effects.
Anthocyanins from cranberries exhibit anti-atherosclerotic functions through multiple mechanisms (Currie et al.,
2022). On the one hand, they inhibit platelet aggregation and the oxidative modification of low-density lipoprotein (LDL), and suppress the abnormal proliferation of smooth muscle cells (Gaiz et al.,
2020). On the other hand, they prevent excessive proliferation of endothelial cells, effectively reducing the risk of atherosclerosis (Ciumărnean et al.,
2020; Garcia and Blesso,
2021). Anthocyanins in wine can reduce the aggregation of low-density lipoprotein in the human body and modulate cell signaling pathways within blood vessels, contributing to vascular health (Zhang et al.,
2021). In vitro experiments have shown that anthocyanins from purple sweet potato can reduce adipogenesis and break down excess lipids, thereby exerting lipid-lowering effects (Ju et al.,
2011). Cyanidin can regulate AMP-activated protein kinase (AMPK) to maintain hepatic lipid homeostasis (Inaguma et al.,
2011). Anthocyanins from black rice can improve insulin sensitivity by modulating the expression of related genes, which may help prevent and ameliorate type II diabetes (Zhao et al.,
2024). For symptoms of visual blurring caused by asthenopia, anthocyanins can effectively alleviate these symptoms and exhibit adjuvant therapeutic effects on diabetic retinopathy (Li et al.,
2025b).
3.4 Anti-inflammatory and antibacterial effects
Inflammation is a non-specific immune response of the body to various stimuli, and chronic inflammation can induce a variety of diseases (Gelon et al.,
2022). Maize anthocyanins and proanthocyanidins exert anti-inflammatory effects by specifically suppressing the activities of key inflammatory enzymes COX-2 and iNOS. Furthermore, these bioactive components effectively block the activation of NF-κB/JNK signaling cascades and downregulate the secretion of pro-inflammatory mediators TNF-α and MCP-1 in both adipocytes and macrophages, thereby alleviating excessive inflammatory responses (Colombo et al.,
2021; Rossi et al.,
2012). Regarding the underlying mechanism, anthocyanins achieve their anti-inflammatory effects by modulating the function of THP-1 cells and activating the MAPK (mitogen-activated protein kinase) signaling pathway (Min et al.,
2010). Anthocyanins exhibit broad-spectrum inhibitory activity against foodborne pathogens, with the mechanism involving disruption of cell membrane integrity and interference with protein homeostasis (Zhou et al.,
2020). Dose-effect studies have shown that anthocyanins can effectively inhibit biofilm formation by
Listeria (Liu et al.,
2022a). Regarding the modulation of the gut microecological environment, anthocyanins promote the growth of
Bifidobacterium and inhibit the adhesion and colonization of
Escherichia coli through nutrient competition (Liu et al.,
2022a).
In view of the considerable health benefits conferred by high-anthocyanin maize—including antioxidative, anti-inflammatory, and vision-protective activities—the development of efficient and robust separation protocols for these bioactive constituents represents a pivotal bottleneck in the transition from laboratory research to functional food manufacturing and industrial-scale processing. Accordingly, the following discussion will systematically delineate the established technological routes for anthocyanin extraction and purification, with particular emphasis on current optimization strategies and emerging trends.
4 Extraction and Purification of Anthocyanins
4.1 Qualitative and quantitative analysis of anthocyanins
Anthocyanins, as important water-soluble pigments in nature, are responsible for the bright red and purple colors of most fruits and vegetables. Purple corn, which is rich in anthocyanins, serves as a natural food colorant (Lu et al.,
2024). In South American countries such as Peru and Bolivia, anthocyanins derived from purple corn have been utilized for centuries (Pu et al.,
2007). Qualitative analysis determines whether anthocyanins are present in a sample, while quantitative analysis measures the concentration of anthocyanins. The technical methods for qualitative and quantitative analysis of anthocyanins include spectrophotometry, chromatography, and mass spectrometry (Fernandez-Aulis et al.,
2019). Spectrophotometry is relatively simple to perform and is commonly employed for routine total content determination and preliminary qualitative analysis. UV-visible spectrophotometry determines the total anthocyanin glycoside content in a solution by measuring the characteristic absorption peaks of anthocyanin molecules at specific wavelengths (Kumkum et al.,
2024). High-performance liquid chromatography (HPLC), by contrast, separates anthocyanin components in a mixture through chromatographic separation, followed by qualitative and quantitative analysis using appropriate detectors (Zhao et al.,
2017; Jin et al.,
2025). Among these, HPLC coupled with a diode array detector (HPLC-DAD) enables simultaneous acquisition of full-wavelength UV-visible spectra after chromatographic separation. It allows quantification based on the characteristic absorption of anthocyanins at approximately 520 nm and preliminary qualitative identification through retention times and spectral features, making it suitable for routine determination of total anthocyanin content and comparative analysis among varieties (Ma et al.,
2024; Saha et al.,
2020). Mass spectrometry, by ionizing and dissociating anthocyanin molecules in the sample, enables the identification of molecular structures, contents, and compositional profiles of anthocyanin glycosides (Fernandez-Aulis et al.,
2019). Liquid chromatography coupled with tandem mass spectrometry (LC-MS/MS) combines the separation capability of chromatography with the structural elucidation capability of mass spectrometry. It enables the determination of anthocyanidin types, glycosyl moieties, and acyl substitutions based on quasi-molecular ions and characteristic fragment ions. In multiple reaction monitoring (MRM) mode, it also allows highly sensitive quantification of trace anthocyanins, making it an essential tool for the identification of individual anthocyanins and their accurate quantification in complex matrices (de Pascual-Teresa et al.,
2002; Abdin et al.,
2020).
4.2 Extraction of anthocyanins
Purple (black) maize possesses advantages such as high anthocyanin content and wide availability of raw materials, making it a suitable feedstock for natural pigment extraction (Fernandez-Aulis et al.,
2019). Based on differences in anthocyanin solubility in an ethanol-water system, solvent permeation can be employed to facilitate the diffusion of active constituents from plant cells into the liquid phase, followed by solvent recovery and product concentration through vacuum distillation (Xue et al.,
2020; Jin et al.,
2025). To prevent the hydrolysis of non-acylated anthocyanins during extraction, an appropriate amount of citric acid or tartaric acid can be added to the extraction solvent as a protective agent (Fernandez-Aulis et al.,
2019). Ultrasound-assisted extraction is a widely used method with relatively high extraction efficiency (Nurkhasanah et al.,
2023; Jovanović et al.,
2020); its principle lies in the cavitation effect, which generates localized high temperature and pressure, causing instantaneous disruption of plant cell structures and accelerating the release of target compounds (Chen et al.,
2022b; Corrales et al.,
2008). The microbial fermentation method degrades the plant cell wall through the action of enzymes or microorganisms, thereby promoting the rapid release of intracellular anthocyanins and enhancing the efficiency of subsequent extraction (Kuntz et al.,
2016).
4.3 Separation and purification of anthocyanins
Three core technologies are primarily relied upon in the industrial purification of anthocyanins: high speed countercurrent chromatography (HSCCC), macroporous resin adsorption, and membrane separation (Fernandez-Aulis et al.,
2019; Liu et al.,
2022b). HSCCC utilizes polytetrafluoroethylene (PTFE) tubing to establish a dynamic partitioning equilibrium in a two-phase system during rotation (Abdin et al.,
2020). This technique features continuous separation capability, simple operation, large processing capacity, and high separation efficiency (Li et al.,
2013). Macroporous resins possess a large specific surface area and strong adsorption capacity and are widely used for the preliminary enrichment of plant anthocyanins (Yue et al.,
2016; Jin et al.,
2025). For example, D3520 resin can substantially increase the purity of blueberry anthocyanins. Membrane separation technology, based on the principle of molecular weight cut-off, achieves fractionation and purification of anthocyanins and offers the advantages of continuous operation, energy efficiency, and preservation of high biological activity (He et al.,
2017).
The efficiency of the above-described extraction and purification processes depends on the consistency and predictability of raw material traits, directing research attention from downstream processing upstream to germplasm improvement. Accordingly, the following discussion will systematically address how breeding progress in high-anthocyanin maize underpins extraction efficiency, with a focus on three aspects: spatiotemporal anthocyanin accumulation patterns, breeding strategies, and the deployment of key biosynthetic genes in genetic improvement.
5 Development of High-Anthocyanin Maize
5.1 Distribution patterns and breeding targets of anthocyanins in maize
In maize, anthocyanins are distributed across multiple organs, including stems, husks, cobs, and tassels, in addition to kernels (Deineka et al.,
2016; Chaiittianan et al.,
2017). The anthocyanin content of individual organs exhibits dynamic temporal variation across developmental stages: leaf content peaks at the large bell mouth stage or at anthesis; stem content increases before declining, with its maximum occurring at the large bell mouth stage; tassel content reaches its highest level at anthesis and subsequently decreases; husk and cob anthocyanins increase steadily from the grain filling stage through to physiological maturity, reaching their maxima at maturity. Despite variation in the timing of peak accumulation among organs, total anthocyanin content at the whole-plant level demonstrates an overall increasing trend throughout the growth cycle (Zhu,
2011). Regarding breeding target selection, the kernel aleurone layer and pericarp represent the principal accumulation sites for edible high-anthocyanin maize, with their contents differing by a factor of 6 to 8 between the two tissues; moreover, these sites are governed by independent MBW regulatory modules (Chatham et al.,
2021). Anthocyanin contents in husks and cobs peak at maturity, and in certain genotypes, husk content can reach ten times that of kernels (Cai et al.,
2023), indicating that organs other than kernels may serve as alternative targets for industrial extraction-oriented high-anthocyanin maize. Elucidating the accumulation patterns across different tissues is a prerequisite for formulating differentiated breeding objectives for edible and extraction-type varieties.
5.2 Breeding approaches for high-anthocyanin maize
5.2.1 Conventional hybrid breeding and marker-assisted selection
Conventional hybrid breeding remains the predominant approach for developing high-anthocyanin maize varieties. Studies have demonstrated that both grain anthocyanin yield and total grain yield exhibit high heritability; however, yield components such as ear diameter and kernel set exert significant effects on total production (Shi et al.,
2012). Research involving 16 hybrid combinations derived from 18 inbred lines revealed substantial variation in anthocyanin content among hybrids, with total anthocyanin yield per unit area determined jointly by grain anthocyanin concentration and grain yield. Notably, grain anthocyanin content shows a highly significant negative correlation with grain yield, whereas leaf anthocyanin content is negatively correlated with photosynthetic rate (Shi et al.,
2012). This antagonistic relationship constitutes a major dilemma in conventional breeding aimed at the simultaneous improvement of anthocyanin content and yield. Crossing the haploid inducer line CAU6 with purple embryo materials to construct a Multi-parent Advanced Generation Inter-Cross (MAGIC) breeding system can reduce the generation cycle of doubled haploid (DH) lines by 40% (Chen et al.,
2022a), thereby accelerating the homozygosity and stabilization of purple inbred lines. Evaluation of 12 fresh-market maize varieties indicated an anthocyanin content range of 17–188 mg/100 g, with Heitiannuo 188 exhibiting the highest level; grain anthocyanin content showed a significant negative correlation with soluble protein content and a highly significant positive correlation with starch content (Jiang et al.,
2025). With the functional characterization of anthocyanin biosynthetic regulatory genes, marker-assisted selection (MAS) based on functional gene variants has emerged as a useful tool for enhancing breeding efficiency. The allelic state of
Pr1 determines whether kernels display red (pelargonidin) or purple (cyanidin) pigmentation, serving as an early screening marker for color classification (Peniche-Pavía et al.,
2022). Allelic variations at
C1,
R1,
B1, and
Pl1 are significantly associated with anthocyanin accumulation, enabling the elimination of off-type individuals at the seedling or seed stage through genotyping, thereby shortening the phenotypic identification cycle (Hichri et al.,
2011). Nevertheless, the current MAS system for maize anthocyanins remains confined to a limited number of classical genes, lacking genome-wide quantitative trait locus (QTL) markers tightly linked to anthocyanin content, which constrains the efficiency of pyramiding minor-effect polygenes.
5.2.2 Transgenic and genome editing technologies
Transgenic technology enables the circumvention of intraspecific genetic barriers and the efficient expression of exogenous or homologous regulatory genes (Yadava et al.,
2017). Researchers at the Biotechnology Research Institute of the Chinese Academy of Agricultural Sciences developed a multigene expression vector system employing an embryo-specific bidirectional promoter coupled with 2A peptide-linked cassettes; introduction of
ZmBz1,
ZmBz2,
ZmC1, and
ZmR2 into maize yielded purple maize lines with anthocyanin enrichment in both the embryo and endosperm, reaching a maximum dry weight content of 291 mg/100 g (Liu et al.,
2018). Furthermore, crossing T
1 transgenic segregants with the inbred line B73 revealed that F
1 progeny displayed purple pigmentation in glumes, husks, leaf sheaths, and brace roots, whereas leaves remained green, silks were predominantly yellow, and kernels were mostly yellow; importantly, days to anthesis and silking, ear height, and plant height remained comparable to those of the control (Feng et al.,
2018). These findings demonstrate that exogenous regulatory gene introduction enables anthocyanin accumulation in specific tissues; however, precise spatial control of expression remains technically challenging. Constitutive overexpression may induce pigmentation in photosynthetic organs, thereby compromising light-use efficiency, or may disturb normal grain coloration. Additionally, introduction of a 35S promoter-driven
A1 cDNA into anthocyanin-deficient mutants restored red pigment synthesis (Haselmair-Gosch et al.,
2018; Kumkum et al.,
2024). Gene editing technology offers novel tools for the precise modification of anthocyanin regulatory networks. Researchers at the Beijing Academy of Agriculture and Forestry Sciences employed base editing to modify
ZmR1, successfully achieving high-level anthocyanin accumulation throughout the entire maize plant (Luo et al.,
2022). BSA-seq analysis of an F
2 population derived from a cross between black and white waxy maize identified the structural gene Zm00001eb276450 and the transcription factor genes Zm00001eb276870 and Zm00001eb276640 as candidate regulators of kernel anthocyanin accumulation. These novel loci, together with previously characterized genes including
ZmC1,
ZmR2,
ZmBz1, and
ZmBz2 represent promising targets for genomic selection breeding of high-anthocyanin maize (Zhang et al.,
2025a).
5.3 Breeding of high-anthocyanin maize varieties (black/purple maize) in China in recent years
The breeding of high-anthocyanin maize cultivars in China has a relatively long history, with fresh-eating maize being the primary focus of these programs (Luo et al.,
2022). Owing to regional differences in consumer preferences for the eating quality of fresh maize between northern and southern China, cultivars developed in northern regions are predominantly waxy type, whereas those bred in southern regions are predominantly sweet type.
In recent years, representative high-anthocyanin maize cultivars developed in northern China include the waxy varieties Jingzinuo 519, Jingzinuo 318, Jingzinuo 219, and Jinghuanuo 2008 (bred by the Maize Research Institute, Beijing Academy of Agriculture and Forestry Sciences); Jinheinuo 1, Jinheinuo 2, and Heitiannuo 636 (Shanxi Agricultural University); Heinuo 808 (College of Agronomy, Xinjiang Agricultural University); Zhengheinuo 8 (Institute of Food Crops, Henan Academy of Agricultural Sciences); Luheitiannuo 201 (Shandong Academy of Agricultural Sciences); and Jinheinuo 529 (Tianjin Agricultural University). In southern China, representative cultivars include the sweet and waxy varieties Ziyujingtian (Sichuan Agricultural University); Zitiannuo 2020 (Chongqing Three Gorges Academy of Agricultural Sciences); Heitian 107 and Yuheitiannuo 950 (Chongqing Academy of Agricultural Sciences); Suyuzinuo 903 and Suyuzinuo 706 (Jiangsu Yanjiang Institute of Agricultural Sciences); Enheitian 18 (Enshi Tujia and Miao Autonomous Prefecture Academy of Agricultural Sciences); Heitiannuo 5201 and Zheheitian 208 (Institute of Maize and Characteristic Dryland Crops, Zhejiang Academy of Agricultural Sciences); Heitiannuo 168 (Zhejiang Dongyang Maize Research Institute); Yueheitiannuo 318 (Institute of Crop Sciences, Guangdong Academy of Agricultural Sciences); Minhuatiannuo 136 and Minzinuo 1 (Institute of Crop Sciences, Fujian Academy of Agricultural Sciences); and Guihuatiannuo 540 and Guihuatiannuo 531 (Guangxi Academy of Agricultural Sciences). For silage maize, Zijing 1 (originally designated Anthocyanin 1), developed by the Institute of Horticulture, Anhui Academy of Agricultural Sciences, is noteworthy. Additionally, Ziyu 194, bred by Shanxi Agricultural University, is a special-purpose cultivar with an anthocyanin content of 0.246 g/100 g (Meng et al.,
2020).
The above-mentioned high-anthocyanin maize cultivars are mostly fresh-eating varieties. Breeding priorities for these materials include plant architecture, ear height, disease resistance, as well as grain appearance and cooking quality. These selection criteria largely follow those established for conventional fresh-eating maize, with no dedicated thresholds set for anthocyanin content. Cultivar data summarized in this section were retrieved from the China Big Data Platform for Seed Industry (Department of Seed Industry Administration, Ministry of Agriculture and Rural Affairs).
5.4 Major challenges in high-anthocyanin maize breeding
Despite increasing diversification in breeding methodologies, the industrialization of high-anthocyanin maize continues to encounter substantial challenges that require integrated solutions through synergistic advances combining basic research and breeding strategy innovation. First, multiple studies have established significant negative correlations between grain anthocyanin content and both grain yield and soluble protein content (Shi et al.,
2012; Jiang et al.,
2025), as well as between leaf anthocyanin content and photosynthetic rate. Consequently, pyramiding of high-anthocyanin alleles through genomic selection alone is insufficient to overcome the yield penalty. Second, anthocyanin accumulation is profoundly influenced by environmental variables including light intensity, temperature, and nitrogen availability, resulting in considerable genotype-by-environment interaction that generates large fluctuations in anthocyanin content across years and locations, thereby constraining variety registration and commercial deployment. Moreover, unlike yield and disease resistance, anthocyanin content has not yet been incorporated as a quantifiable quality parameter in official variety approval standards, depriving breeders of positive incentives to prioritize this trait as a core selection objective.
Collectively, these breeding advances have generated extensive high-anthocyanin maize germplasm, establishing a critical resource base for industrial exploitation; nevertheless, commercial translation remains subject to numerous practical constraints, which are examined in the following sections through the lenses of utilization pathways, existing bottlenecks, and market potential.
6 Development of the High-Anthocyanin Maize Industry
6.1 Applications and utilization of high-anthocyanin maize
High-anthocyanin maize is rich in natural antioxidant compounds and holds considerable promise for agricultural production and the food industry (Cai et al.,
2023). Based on end-use applications, high-anthocyanin maize can currently be categorized into five types: (1) Fresh-market maize. This category represents the most commercially advanced and can be promoted through multi-site trials following fresh maize variety approval standards. However, current national fresh maize approval standards lack mandatory anthocyanin content specifications, resulting in considerable variation in color intensity and anthocyanin levels among black maize products on the market. Additionally, anthocyanins are prone to degradation during postharvest storage of fresh maize. How to maintain the characteristic tender texture and flavor of fresh maize while enhancing anthocyanin content remains a key challenge requiring coordinated optimization between breeding and processing. (2) Dry-grain maize. Its potential applications include functional staple food, food ingredients (e.g., black maize flour, black maize flakes), and premium livestock feed. However, this category remains at a preliminary research stage, with neither systematic quality evaluation systems nor formal variety approval pathways established. During harvesting, threshing, drying, and long-term storage, anthocyanins undergo significant thermal and oxidative degradation. Existing drying equipment and storage facilities are designed for conventional maize, lacking technical specifications for low-temperature drying and light-protected storage aimed at anthocyanin preservation. (3) Silage maize. This type emphasizes whole-plant utilization. Although no officially approved varieties currently exist, certain lines have entered regional production. During ensiling, fermentation temperature, pH, and microbial communities may significantly influence anthocyanin stability. However, existing silage quality evaluation systems do not incorporate anthocyanin content or antioxidant activity parameters, precluding quantitative assessment of the nutritional value of high-anthocyanin silage maize. (4) Anthocyanin extraction-dedicated maize. Currently, no specialized extraction varieties exist domestically. Raw materials are mostly sourced from fresh-market or dry-grain varieties, resulting in substantial fluctuations in raw material anthocyanin content and pronounced supply seasonality that cannot meet the demands of continuous industrial production. Furthermore, high-value utilization pathways for by-products such as extracted corn residues and husks remain undeveloped, constraining the economic feasibility of whole-component utilization. (5) Multi-functional dual-purpose maize. This category balances grain yield with plant biomass, enabling dual use for grain and feed or combined grain production and extraction, and is regarded as a promising type for future development. However, the "multi-functional" positioning itself introduces ambiguity in breeding objectives: if grain consumption is prioritized, anthocyanins must be concentrated in the aleurone layer with moderate kernel hardness; if whole-plant silage or extraction is prioritized, anthocyanin content in stems and husks must be enhanced alongside biomass. The divergent requirements for anthocyanin tissue distribution, content thresholds, and agronomic traits across different uses make current breeding systems insufficient, as they lack integrated selection indices tailored for multi-functional objectives.
6.2 Critical constraints and strategic countermeasures for high-anthocyanin maize industrialization
To bridge the gap from cultivar development to market commercialization, the high-anthocyanin maize industry must overcome deep-seated constraints spanning standardization, processing technology, and market dynamics, extending beyond continuous genetic improvement. (1) Industrial disorder stemming from absent standardization frameworks. The sector currently lacks unified sampling and analytical protocols; disparate methodologies and reporting units across research groups and enterprises generate inconsistent data among varieties and batches, thereby distorting market valuation. (2) Insufficient deep-processing capacity limiting value capture. Value-added processing remains rudimentary, with notable delays in high-purity anthocyanin monomer isolation, stabilized formulation development, and systematic functional assessment. (3) Divergence between market consumption patterns and product positioning. While health-conscious consumer segments may demonstrate receptivity toward premium-priced high-anthocyanin maize, mainstream market expansion could represent a potential strategy warranting evaluation through production scaling and enhanced awareness of anthocyanin-mediated health benefits.
Interventions addressing these constraints should focus on three strategic dimensions. First, establish a mandatory, tiered classification standard system. Relevant authorities are advised to promptly formulate industry standards for high-anthocyanin maize, incorporating anthocyanin content into quality assessment protocols for fresh-market maize variety approval. Second, advance low-temperature processing technologies. For dry grain, develop low-temperature circulating drying and nitrogen-flushed, light-protected storage technologies to substantially reduce maize anthocyanin degradation; for fresh-market maize, promote rapid post-harvest pre-cooling (0−4 °C) to extend shelf life (Zhang et al.,
2023). Third, construct an integrated "variety, production base, brand" industrial chain. Leveraging existing high-anthocyanin varieties, establish specialized production bases in ecologically suitable regions, operating under contract farming models characterized by unified seed supply, cultivation protocols, harvesting standards, and brand marketing. Deploy blockchain traceability technology to record variety pedigrees, anthocyanin assay certificates, and plot-level agronomic data on a distributed ledger, thereby addressing consumer trust issues. For the fresh-food market, develop convenient products such as ready-to-eat black maize segments and black maize pulp packs to lower consumption barriers. For the extraction-oriented market, enter into long-term raw material supply agreements with functional food enterprises to lock in cultivation area and procurement prices, thereby mitigating seasonal volatility risks in raw material supply.
6.3 Application potential and market prospects of high-anthocyanin maize
The application potential of high-anthocyanin maize must be assessed within the dual context of the global anthocyanin industry and the Chinese fresh maize market to avoid speculative generalizations divorced from industrial realities. Globally, natural anthocyanins, serving as food colorants and functional ingredients, are experiencing steady growth in annual demand. Compared with conventional sources such as blueberries and purple sweet potatoes, maize offers distinct agronomic advantages, including high yield potential, high mechanization compatibility, and low production costs. Domestically, China is home to over 500 fresh corn processing enterprises, with a total output value exceeding 25 billion yuan, ranking first globally in industrial scale (He et al.,
2026). In the fresh food segment, the breakthrough strategy for high-anthocyanin maize centers on functional visualization, which entails explicit disclosure of anthocyanin content on packaging and quantitative benchmarking against competing products such as blueberries to establish consumer value perception. In the dry-grain segment, staple-oriented products such as black maize flour, black maize flakes, and black maize meal replacement powders can be developed for the functional cereal market (Verediano et al.,
2022). For silage applications, the "antioxidant silage" concept can be promoted to high-end dairy operations, leveraging improved milk quality as a value proposition to support feed price premiums (Zhang et al.,
2025b). Integrated whole-plant resource development represents another important lever for enhancing industrial economic returns. In certain varieties, husks exhibit high anthocyanin content coupled with loose fibrous architecture, rendering them suitable feedstock for anthocyanin extraction. Following extraction, cobs can be processed into activated carbon or biomass substrates, while residual anthocyanins in ensiled stalks retain antioxidant activity capable of improving ruminant intestinal health (Yang et al.,
2021; Xu et al.,
2026). Through a cascading utilization model encompassing "grain consumption, husk extraction, stalk silage, and cob valorization," the output value per unit area can be substantially elevated, achieving genuine synergy among economic, ecological, and social benefits.
The high-anthocyanin maize industry has reached a critical inflection point, transitioning from a purely variety-driven paradigm toward one shaped by standardization, technological innovation, and market forces. Overcoming bottlenecks across the entire value chain, including variety registration, standard development, processing technology, and brand establishment, is essential for translating laboratory-derived, high-anthocyanin germplasm into competitive field-scale industries, thereby advancing national health objectives and the high-quality development of agriculture.
7 Perspectives
With the sustained expansion of China’s economy and continuous improvement in residents’ living standards, consumer demand for high-quality agricultural products has been steadily rising. High-anthocyanin maize well conforms to public demands for nutritional wellness and possesses considerable domestic market potential. At the molecular level, although the core biosynthetic pathway of anthocyanin and the MBW regulatory framework in maize have been largely clarified, the fine-scale regulatory networks underlying tissue-specific anthocyanin accumulation remain poorly understood. Current research has predominantly focused on anthocyanin synthesis in the aleurone layer and pericarp, whereas the genetic basis for efficient anthocyanin accumulation in non-kernel organs—including husks, cobs, and stalks—remains largely uncharacterized. Future efforts should integrate single-cell transcriptomics, spatial metabolomics, and epigenetic approaches to generate spatiotemporal atlases of regulatory factor expression across tissues and developmental stages, thereby achieving a deeper dissection of the molecular and genetic mechanisms governing anthocyanin biosynthesis in maize. Regarding health-promoting functionality, existing studies have established that maize anthocyanins exhibit antioxidant, anti-inflammatory, and cardioprotective activities, and specific extracts have also shown promising potential for cancer chemoprevention. Nevertheless, the systemic metabolic pathways underlying these bioactive effects and their clinical translational value require further systematic evaluation and verification. From a breeding perspective, resolving the genetic trade-off between high anthocyanin content and yield penalty represents a core scientific challenge. To address this issue, CRISPR/Cas9-mediated base editing technology can be applied to conduct saturation editing on key loci associated with anthocyanin synthesis, aiming to enhance pigment accumulation without compromising photosynthetic efficiency or grain filling. At the industrial application level, the absence of unified and standardized quality criteria constitute a critical bottleneck constraining market commercialization. Moving forward, anthocyanin content and color stability should be incorporated into quality evaluation systems for variety approval. Meanwhile, distinct categories including fresh market, dry grain, silage, extraction-dedicated, and multifunctional dual-use types should be clearly defined to advance standardized industrial development.
In summary, future research on high-anthocyanin maize needs to break through the limitations of single-trait improvement and pivot toward integrated innovation across the entire value chain encompassing synthetic mechanisms, genetic regulation, environmental response, processing stability, and market standardization. The integration of precise molecular-design breeding with robust industrial standardization systems will facilitate the transformation of high-anthocyanin maize from laboratory-specific germplasm resources to competitively advantageous industrial products. Ultimately, this strategy will support the dual objectives of enhancing national nutritional health and advancing high-quality agricultural development.
High-Anthocyanin maize: From biosynthesis to industrial applications
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