Table 1.
A comprehensive list of cold acclimatization genes, transcription factors, and proteins, along with the species of identification, their mode of action, and references, is provided.
| Gene/Transcription Factor/Protein | Species | Mode of Action | References |
|---|---|---|---|
| Fruit Trees and Shrubs | |||
| MdCPK1a | Malus domestica (apple) | Overexpression in tobacco increased freezing, salt, and cold tolerance, root length, and antioxidants, while reducing membrane damage and lipid peroxidation | (Dong et al., 2020) |
| PpCBF6 | Prunus persica (peach) | Transient overexpression prevented sucrose degradation and increased chilling tolerance in peach fruit. | (Cao et al., 2021) |
| PpCBF1 | Overexpression in apple increased freezing tolerance in both non-acclimated and acclimated conditions | (Wisniewski et al., 2011) | |
| PpyCBF1-3 | Pyrus pyrifolia (Asian pear) | Overexpression in Arabidopsis enhanced tolerance to low temperature, salt, and drought stresses while reducing ROS production | (Ahmad et al., 2019) |
| BB-CBF | Vaccinium corymbosum (Highbush blueberry) | Overexpression in Arabidopsis enhanced freezing tolerance and induced certain COR genes | (Polashock et al., 2010) |
| VvCBF2 | Vitis vinifera (grape) | Overexpression in Arabidopsis enhanced cold tolerance | (Takuhara et al 2011) |
| VvCBF4 | |||
| VvCBFL | |||
| VvCZFPL | |||
| PgCBF3 | Punica granatum (pomegranate) | Overexpression in Arabidopsis increased cold resilience by raising proline, total soluble sugar levels, and enzymatic activity (CAT, SOD, and peroxidase), while reducing electrolyte leakage, MDA content, and ROS production | (Wang et al., 2022) |
| PgCBF7 | |||
| PtCBF | Poncirus trifoliata (trifoliate orange) | Exhibited increased accumulation in response to low temperature | (He et al., 2012) |
| MaPIP2-7 | Musa acuminata L. (banana) | Overexpression enhanced tolerance to multiple abiotic stresses (low temperature, drought, salt) by maintaining osmotic balance, reducing membrane injury, and increasing chlorophyll, proline, soluble sugar, and ABA | (Xu et al., 2020) |
| MusaPIP1;2 | Overexpression improved cold resilience by reducing MDA levels, increasing proline and relative water content, and enhancing photosynthetic efficiency | (Sreedharan et al., 2013) | |
| DlCBF1-3 | Dimocarpus longan (longan) | Overexpression in Arabidopsis improved cold tolerance by increasing proline accumulation, reducing ROS content, and upregulating cold-responsive genes in the CBF pathway | (Yang et al., 2020) |
| Solanum | |||
| LeCOR413PM2 | Lycopersicon esculentum (tomato) | Overexpression prevented membrane damage by increasing antioxidant enzymes, ROS scavenging, reducing PSII photoinhibition, and improving osmotic regulation, while suppressing by RNAi resulted in increased sensitivity of plans to cold | (Zhang et al., 2021) |
| LeGPA1 | Overexpression increased cold tolerance by inducing ICE-CBF pathway genes and enhancing SOD, peroxidase, CAT, proline, and total soluble sugar levels, while reducing ROS production and lipid peroxidation | (Guo et al., 2020) | |
| AtCBF1 | Heterology expression of the Arabidopsis AtCBF1 in tomato improved plant resilience to low temperature by inducing CAT1 gene expression and reducing H2O2 levels. It also enhanced tolerance to oxidative damage from methyl viologen | (Hsieh et al., 2002) | |
| AtCBF1-3 | Solanum tuberosum (potato) | Overexpression of Arabidopsis AtCBF genes resulted in improved freezing tolerance in potato | (Pino et al., 2007) |
| Field Crops | |||
| ZmDREB1A | Zea mays (maize) | Overexpression in Arabidopsis induced COR genes and conferred plant’s tolerance to cold, drought and high salinity stresses | (Qin et al., 2004) |
| OsDREB1A | Oryza sativa (rice) | Overexpression in Arabidopsis conferred plant’s tolerance to cold, drought and high salinity stresses | (Dubouzet et al., 2003) |
| OsDREB1A/B | Overexpression in Arabidopsis improved tolerance to drought, high-salt and low-temperature. Also, it increased the contents of osmoprotectants such as free proline and various soluble sugars | (Ito et al., 2006) | |
| GmDREB1B | Glycine max (soybean) | Overexpression in Arabidopsis resulted in elevated tolerance to abiotic and environmental stresses such as cold, drought, salinity, and heat | (Kidokoro et al., 2015) |
| BaAFP-1 | Hordeum vulgare (malting barley) | Increased accumulation observed after cold acclimatization | (Ding et al., 2015) |
| IbCBF3 | Ipomoea batatas (sweet potato) | Overexpression led to enhanced tolerance to cold, drought and oxidative stress, and showed improved photosynthesis efficiency and reduced hydrogen peroxide levels | (Jin et al., 2017) |
| BnCBF5, BnCBF 17 | Brassica napus (rapeseed) | Overexpression improved freezing tolerance, COR genes mRNA accumulation, photosynthesis-related gene transcript levels accumulation, and chloroplast development resulting in increased photosynthetic efficiency and capacity | (Savitch et al., 2005) |
| MfLEA3 | Medicago falcata (yellow alfalfa) | MfLEA3 was induced by cold, dehydration, and ABA. Its constitutive expression enhanced tolerance to cold, drought, and high-light stress in transgenic tobacco plants, along with higher CAT activity. | (Shi et al., 2020) |
| Ornamental Plants | |||
| SikCOR413PM1 | Saussurea involucrata (snow lotus) | Overexpression in tobacco enhanced cold tolerance through Ca2+ signaling and membrane stabilization | (Guo, et al., 2019a) |
| SiDHN | Overexpression in tomato enhanced cold tolerance by preserving cell membrane integrity, increasing chlorophyll a and b contents, carotenoid, reducing chlorophyll photo-oxidation and ROS accumulation, and improving antioxidant enzyme activity and photochemical electron transfer efficiency | (Guo, et al., 2019b) | |
| Herbaceous Plants | |||
| AtCBF2 | Arabidopsis thaliana | Loss-of-function mutants resulted sensitivity to freezing after cold acclimatization and high salinity | (Zhao et al., 2016) |
| OST1 | Overexpression enhanced freezing tolerance, while ost1 mutants showed freezing hypersensitivity. Cold-activated OST1 phosphorylates ICE1 and boosts its stability and transcriptional activity in the CBF pathway. | (Ding et al., 2018) | |
| Phospholipase Dδ (PLDδ) | Overexpression increased freezing tolerance, while knockout increased sensitivity. PLDδ gene is involved in membrane lipid hydrolysis, contributing ~20% to phosphatidic acid production; its overexpression enhanced phosphatidic acid production | (Li et al., 2004) | |
| AtDREB1A | Nicotiana tabacum (tobacco) | Overexpression of Arabidopsis AtDREB1A gene enhanced drought and cold stress tolerance in tobacco by inducing abiotic stress-related genes and interacting with the dehydration responsive element | (Kasuga et al., 2004) |
| PpCBF3 | Poa pratensis (kentucky bluegrass) | Transient overexpression in Arabidopsis increased freezing tolerance by reducing electrolyte leakage, H2O2 and O2•− contents, increasing chlorophyll content and photochemical efficacy, and upregulating cold tolerance genes | (Zhuang et al., 2015) |
| BdIRI-7 | Brachypodium distachyon (Purple false brome) | Knockdown mutants exhibited reduced freezing survival and impaired ice-crystal growth restriction in plants, while showing increased membrane damage and electrolyte leakage. | (Bredow et al., 2016) |
| PsCOR413PM2 | Phlox subulate (creeping phlox) | Overexpression in Arabidopsis improved low-temperature tolerance by modulating Ca2+ flux and influencing the expression of stress-related COR and CBF genes | (Zhou et al., 2018) |
| Others | |||
| PttLHY1, PttLHY2 | Populus tremula × Populus tremuloides (poplar) | RNAi knockdown compromised freezing tolerance during winter dormancy | (IbÁñez et al., 2010) |
| EgCBF3 | Elaeis guineensis var. Dura × Pisisfera (oil palm) | Overexpression in tomato improved abiotic stress tolerance under in vitro conditions | (Ebrahimi et al., 2016) |