Kelly Balfour
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Previous Organizations / Institutes
The organizations / institutes from the scholar's other articles in the past ten years
Research Subjects
Kelly Balfour has a total of 44 co-authors. Here are the top co-authors by number of shared publications:
- Hua-Feng Wang
Sanya Nanfan Research Institute of Hainan University, Hainan Yazhou Bay Seed Laboratory, Sanya 572025, China
4 shared publications - Mir Muhammad Nizamani
Guangdong Provincial Key Laboratory of Marine Disaster Prediction and Prevention, Institute of Marine Sciences, Shantou University, Shantou 515063, China
4 shared publications - Salman Qureshi
Institute of Geography (Landscape Ecology), Humboldt University of Berlin, Rudower Chaussee 16, 12489 Berlin, Germany
3 shared publications - Zhi-Xin Zhu
Hainan Key Laboratory for Sustainable Utilization of Tropical Bioresources, College of Tropical Crops , Hainan University, Haikou, China
3 shared publications - Chi Yung Jim
Department of Social Sciences and Policy Studies Education University of Hong Kong Hong Kong China
2 shared publications Gillian Piggott
Department of Biology Queen’s University Kingston Ontario Canada
2 shared publications
Latest Publications
- Open Access
Metabolic Profiling and Antioxidant Analysis for the Juvenile Red Fading Leaves of Sweetpotato
Leaves of sweetpotato (Ipomoea batatas L.) are promising healthy leafy vegetable. Juvenile red fading (JRF) leaves of sweetpotato, with anthocyanins in young leaves, are good candidates for developing functional vegetables. Here, metabolic profiling and possible antioxidants were analyzed for five leaf stages of the sweetpotato cultivar “Chuanshan Zi”. The contents of anthocyanins, total phenolics, and flavonoids all declined during leaf maturation, corresponding to declining antioxidant activities. By widely targeted metabolomics, we characterized 449 metabolites belonging to 23 classes. A total of 193 secondary metabolites were identified, including 82 simple phenols, 85 flavonoids, 18 alkaloids, and eight terpenes. Analysis of the metabolic data indicates that the antioxidant capacity of sweetpotato leaves is the combined result of anthocyanins and many other colorless compounds. Increased levels of “chlorogenic acid methyl ester”, a compromised form of chlorogenic acid, significantly correlated with the declined antioxidant abilities. Besides anthocyanins, some significant metabolites contributing to the high antioxidant property of the sweetpotato leaves were highlighted, including chlorogenic acids, isorhamnetin glycosides, trans-4-hydroxycinnamic acid methyl ester, 4-methoxycinnamic acid, esculetin, caffeate, and trigonelline. This study provides metabolic data for the utilization of sweetpotato leaves as food sources, and sheds light on the metabolomic change for JRF leaves of other plants.
Open AccessMetabolic Profiling and Antioxidant Analysis for the Juvenile Red Fading Leaves of Sweetpotato
Leaves of sweetpotato (Ipomoea batatas L.) are promising healthy leafy vegetable. Juvenile red fading (JRF) leaves of sweetpotato, with anthocyanins in young leaves, are good candidates for developing functional vegetables. Here, metabolic profiling and possible antioxidants were analyzed for five leaf stages of the sweetpotato cultivar “Chuanshan Zi”. The contents of anthocyanins, total phenolics, and flavonoids all declined during leaf maturation, corresponding to declining antioxidant activities. By widely targeted metabolomics, we characterized 449 metabolites belonging to 23 classes. A total of 193 secondary metabolites were identified, including 82 simple phenols, 85 flavonoids, 18 alkaloids, and eight terpenes. Analysis of the metabolic data indicates that the antioxidant capacity of sweetpotato leaves is the combined result of anthocyanins and many other colorless compounds. Increased levels of “chlorogenic acid methyl ester”, a compromised form of chlorogenic acid, significantly correlated with the declined antioxidant abilities. Besides anthocyanins, some significant metabolites contributing to the high antioxidant property of the sweetpotato leaves were highlighted, including chlorogenic acids, isorhamnetin glycosides, trans-4-hydroxycinnamic acid methyl ester, 4-methoxycinnamic acid, esculetin, caffeate, and trigonelline. This study provides metabolic data for the utilization of sweetpotato leaves as food sources, and sheds light on the metabolomic change for JRF leaves of other plants.
- Open Access
- 22 June 2022
- Published by Wiley in Ecology and Evolution
- Vol. 12 (6) , e9006
Smaller species experience mild adversity under shading in an old‐field plant community
Plant competition experiments commonly suggest that larger species have an advantage, primarily in terms of light acquisition. However, within crowded natural vegetation, where competition evidently impacts fitness, most resident species are relatively small. It remains unclear, therefore, whether the size advantage observed in controlled experiments is normally realized in habitats where competition is most intense. We characterized the light environment and tested for evidence of a size advantage in competition for light in an old‐field plant community composed of perennial herbaceous species. We investigated whether larger species contributed to reduced light penetration (i.e., greater shading), and examined the impact of shade on smaller species by testing whether their abundance and richness were lower in plots with less light penetration. Light penetration in plots ranged from 0.3% to 72.4%. Significant effects were more common when analyses focused on small plants that reached reproduction (i.e., flowering rooted units); focusing on only flowering plants (i.e., excluding nonflowering rooted units) can clarify community patterns. Plots with a greater mean species height had significantly lower light penetration, and plots with lower light penetration had significantly lower flowering abundance and richness of small species. However, the impact of shade on the flowering abundance and richness of small species was relatively small (R2 values between 8% and 15%) and depended on how we defined “small species.”Synthesis: Our results confirm that light penetration in herbaceous vegetation can be comparable to levels seen in forests, that plots with taller species cast more shade, and that flowering smaller species are less abundant and diverse in plots where light penetration is low. However, variation in mean plot height explained less than 10% of variation in light penetration, and light penetration explained between 5 and 15% of variation in the flowering abundance and richness of small species. Coupled with the fact that flowering small species were present even within the most heavily shaded plots, our results suggest that any advantage in light competition by large species is limited. One explanation is that at least some small species in these communities are shade‐tolerant. Shade tolerance in predominantly herbaceous communities, particularly among small plant species, requires further research.
- 22 June 2022
- Published by Wiley in Ecology and Evolution
- Vol. 12 (6) , e9006
Open AccessSmaller species experience mild adversity under shading in an old‐field plant community
Plant competition experiments commonly suggest that larger species have an advantage, primarily in terms of light acquisition. However, within crowded natural vegetation, where competition evidently impacts fitness, most resident species are relatively small. It remains unclear, therefore, whether the size advantage observed in controlled experiments is normally realized in habitats where competition is most intense. We characterized the light environment and tested for evidence of a size advantage in competition for light in an old‐field plant community composed of perennial herbaceous species. We investigated whether larger species contributed to reduced light penetration (i.e., greater shading), and examined the impact of shade on smaller species by testing whether their abundance and richness were lower in plots with less light penetration. Light penetration in plots ranged from 0.3% to 72.4%. Significant effects were more common when analyses focused on small plants that reached reproduction (i.e., flowering rooted units); focusing on only flowering plants (i.e., excluding nonflowering rooted units) can clarify community patterns. Plots with a greater mean species height had significantly lower light penetration, and plots with lower light penetration had significantly lower flowering abundance and richness of small species. However, the impact of shade on the flowering abundance and richness of small species was relatively small (R2 values between 8% and 15%) and depended on how we defined “small species.”Synthesis: Our results confirm that light penetration in herbaceous vegetation can be comparable to levels seen in forests, that plots with taller species cast more shade, and that flowering smaller species are less abundant and diverse in plots where light penetration is low. However, variation in mean plot height explained less than 10% of variation in light penetration, and light penetration explained between 5 and 15% of variation in the flowering abundance and richness of small species. Coupled with the fact that flowering small species were present even within the most heavily shaded plots, our results suggest that any advantage in light competition by large species is limited. One explanation is that at least some small species in these communities are shade‐tolerant. Shade tolerance in predominantly herbaceous communities, particularly among small plant species, requires further research.
- Open Access
- 3 March 2022
- Published by Frontiers Media SA in Frontiers in Ecology and Evolution
- Vol. 10
Socio-Ecological Effects on the Patterns of Non-native Plant Distributions on Hainan Island
Non-native plants spread to recipient areas via natural or human-mediated modes of dispersal, and, if the non-native species are invasive, introduction potentially causes impacts on native plants and local ecosystems as well as economic losses. Therefore, we studied the diversity and distributional patterns of non-native plant species diversity in the tropical island province of Hainan, China and its relationships with environmental and socioeconomic factors by generating a checklist of species and subsequently performing an analysis of phylogenetic diversity. To generate the checklist, we began with the available, relevant literature representing 19 administrative units of Hainan and determined the casual, naturalized, or invasive status of each species by conducting field surveys within 14 administrative units. We found that non-native plants of Hainan comprise 77 casual species, 42 naturalized species, and 63 invasive species. Moreover, we found that non-native plant species had diverse origins from North and South America, Africa, and Asia and that the most common species across administrative areas belong to the plant families Asteraceae and Fabaceae. Moreover, the numbers of non-native species distributed in the areas of Hainan bording the coast arer greater than those within interior areas of the province. Among the coastal areas, Haikou has the highest species richness and, simultaneously, the highest values for significantly, positively correlated predictor variables, population and GDP (R2 = 0.60, P < 0.01; R2 = 0.64, P < 0.01, respectively). In contrast, the landlocked administrative units of Tunchang and Ding’an have the smallest number of non-native species, while their populations are less than a quarter of that of Haikou and their GDP less than one tenth. Among natural environmental variables, we determined that the number of non-native species had the strongest correlation with the minimum temperature in the coldest month, which predicts a smaller number of non-native species. Additionally, non-native species are primarily distributed in urban and rural built-up areas and agricultural areas; areas that are dominated by human activities. Overall, our study provides a working checklist of the non-native plants of Hainan as well as a theoretical framework and reference for the control of invasive plants of the province.
- 3 March 2022
- Published by Frontiers Media SA in Frontiers in Ecology and Evolution
- Vol. 10
Open AccessSocio-Ecological Effects on the Patterns of Non-native Plant Distributions on Hainan Island
Non-native plants spread to recipient areas via natural or human-mediated modes of dispersal, and, if the non-native species are invasive, introduction potentially causes impacts on native plants and local ecosystems as well as economic losses. Therefore, we studied the diversity and distributional patterns of non-native plant species diversity in the tropical island province of Hainan, China and its relationships with environmental and socioeconomic factors by generating a checklist of species and subsequently performing an analysis of phylogenetic diversity. To generate the checklist, we began with the available, relevant literature representing 19 administrative units of Hainan and determined the casual, naturalized, or invasive status of each species by conducting field surveys within 14 administrative units. We found that non-native plants of Hainan comprise 77 casual species, 42 naturalized species, and 63 invasive species. Moreover, we found that non-native plant species had diverse origins from North and South America, Africa, and Asia and that the most common species across administrative areas belong to the plant families Asteraceae and Fabaceae. Moreover, the numbers of non-native species distributed in the areas of Hainan bording the coast arer greater than those within interior areas of the province. Among the coastal areas, Haikou has the highest species richness and, simultaneously, the highest values for significantly, positively correlated predictor variables, population and GDP (R2 = 0.60, P < 0.01; R2 = 0.64, P < 0.01, respectively). In contrast, the landlocked administrative units of Tunchang and Ding’an have the smallest number of non-native species, while their populations are less than a quarter of that of Haikou and their GDP less than one tenth. Among natural environmental variables, we determined that the number of non-native species had the strongest correlation with the minimum temperature in the coldest month, which predicts a smaller number of non-native species. Additionally, non-native species are primarily distributed in urban and rural built-up areas and agricultural areas; areas that are dominated by human activities. Overall, our study provides a working checklist of the non-native plants of Hainan as well as a theoretical framework and reference for the control of invasive plants of the province.
- 1 January 2022
- Published by Elsevier in Urban Forestry & Urban Greening
- Vol. 67
Drivers of spontaneous and cultivated species diversity in the tropical city of Zhanjiang, China
- 1 January 2022
- Published by Elsevier in Urban Forestry & Urban Greening
- Vol. 67
Drivers of spontaneous and cultivated species diversity in the tropical city of Zhanjiang, China
- 28 September 2021
- Published by Springer Nature in Urban Ecosystems
- Vol. 25 (2) , 511-522
Response of urban tree DBH to fast urbanization: Case of coastal Zhanjiang in south China
- 28 September 2021
- Published by Springer Nature in Urban Ecosystems
- Vol. 25 (2) , 511-522
Response of urban tree DBH to fast urbanization: Case of coastal Zhanjiang in south China
- Open Access
- 25 September 2021
- Published by Wiley
Smaller species experience mild adversity under shading in an old-field plant community
Plant competition experiments commonly suggest that larger species have an advantage, especially in light acquisition. However, within crowded natural vegetation, where competition evidently impacts fitness, most resident species are relatively small. It remains unclear, therefore, whether the size-advantage observed in controlled experiments is realized in habitats under intensive competition. We tested for evidence of a size-advantage in competition for light in an old-field plant community composed of herbaceous perennial species. We investigated whether larger species contributed to reduced light penetration (i.e., greater shading), and examined the impact of shade on smaller species by testing whether their abundance and richness were lower in plots with less light penetration. Light penetration in plots ranged from 0.3-72.4%. Plots with greater mean species height had significantly lower light penetration. Plots with lower light penetration had significantly lower small species abundance and richness. However, the impact of shade on small species abundance and richness was relatively small (R 2 values between 8% and 15%) and depended on how we defined “small species”. Significant effects were more common when analyses focused on individuals that reached reproduction; focusing on only flowering plants can clarify patterns. Our results confirm that light penetration in herbaceous vegetation can be comparable to levels seen in forests, that plots with taller species cast more shade, and that smaller species are less abundant and diverse in plots where light penetration is low. However, variation in mean plot height explained less than 10% of variation in light penetration, and light penetration explained 5-15% of variation in small species abundance and richness. Coupled with the fact that reproductive small species were present even within the most heavily shaded plots, our results suggest that any advantage in light competition by large species is limited. One explanation is that some small species in these communities are shade tolerant.
- 25 September 2021
- Published by Wiley
Open AccessSmaller species experience mild adversity under shading in an old-field plant community
Plant competition experiments commonly suggest that larger species have an advantage, especially in light acquisition. However, within crowded natural vegetation, where competition evidently impacts fitness, most resident species are relatively small. It remains unclear, therefore, whether the size-advantage observed in controlled experiments is realized in habitats under intensive competition. We tested for evidence of a size-advantage in competition for light in an old-field plant community composed of herbaceous perennial species. We investigated whether larger species contributed to reduced light penetration (i.e., greater shading), and examined the impact of shade on smaller species by testing whether their abundance and richness were lower in plots with less light penetration. Light penetration in plots ranged from 0.3-72.4%. Plots with greater mean species height had significantly lower light penetration. Plots with lower light penetration had significantly lower small species abundance and richness. However, the impact of shade on small species abundance and richness was relatively small (R 2 values between 8% and 15%) and depended on how we defined “small species”. Significant effects were more common when analyses focused on individuals that reached reproduction; focusing on only flowering plants can clarify patterns. Our results confirm that light penetration in herbaceous vegetation can be comparable to levels seen in forests, that plots with taller species cast more shade, and that smaller species are less abundant and diverse in plots where light penetration is low. However, variation in mean plot height explained less than 10% of variation in light penetration, and light penetration explained 5-15% of variation in small species abundance and richness. Coupled with the fact that reproductive small species were present even within the most heavily shaded plots, our results suggest that any advantage in light competition by large species is limited. One explanation is that some small species in these communities are shade tolerant.
- Open Access
- 22 October 2020
- Published by MDPI AG in Remote Sensing
- Vol. 12 (21) , 3477
Using SPOT Data and FRAGSTAS to Analyze the Relationship between Plant Diversity and Green Space Landscape Patterns in the Tropical Coastal City of Zhanjiang, China
Urban green spaces provide a host of ecosystem services, the quantity and structure of which play an important role in human well-being. Rapid urbanization may modify urban green spaces, having various effects on plant diversity. Tropical coastal cities have urbanized rapidly in recent decades, but few studies have been conducted with a focus on their green spaces. We studied the responses of cultivated and spontaneous plants, both key components of urban flora, to the landscape structure of urban green spaces and possible social drivers. We analyzed existing relationships between plant diversity indices, urban green space landscape metrics (using Systeme Probatoire d’Observation de la Terre (SPOT) data,), and social factors, including the type, population density, construction age, and GPS coordinates of each Urban Functional Unit, or UFU. We found that UFUs with more green space patches had higher cultivated and spontaneous species richness than those with fewer green space patches. Spontaneous species richness decreased when green space patches became fragmented, and it increased when green space patches were more connected (e.g., via land bridges). Conversely, cultivated species richness increased with green space patch fragmentation. The phylogenetic diversity of both cultivated and spontaneous plants were weakly associated with green space structure, which was strongly driven by land use. Old UFUs and those with larger populations had more green space patches overall, although they tended to be small and fragmented. Green space patch density was found to increase as the UFU age increased. From the viewpoint of knowledge transfer, understanding the effects and drivers of landscape patterns of urban green spaces could inform the development of improved policies and management of urban green space areas.
- 22 October 2020
- Published by MDPI AG in Remote Sensing
- Vol. 12 (21) , 3477
Open AccessUsing SPOT Data and FRAGSTAS to Analyze the Relationship between Plant Diversity and Green Space Landscape Patterns in the Tropical Coastal City of Zhanjiang, China
Urban green spaces provide a host of ecosystem services, the quantity and structure of which play an important role in human well-being. Rapid urbanization may modify urban green spaces, having various effects on plant diversity. Tropical coastal cities have urbanized rapidly in recent decades, but few studies have been conducted with a focus on their green spaces. We studied the responses of cultivated and spontaneous plants, both key components of urban flora, to the landscape structure of urban green spaces and possible social drivers. We analyzed existing relationships between plant diversity indices, urban green space landscape metrics (using Systeme Probatoire d’Observation de la Terre (SPOT) data,), and social factors, including the type, population density, construction age, and GPS coordinates of each Urban Functional Unit, or UFU. We found that UFUs with more green space patches had higher cultivated and spontaneous species richness than those with fewer green space patches. Spontaneous species richness decreased when green space patches became fragmented, and it increased when green space patches were more connected (e.g., via land bridges). Conversely, cultivated species richness increased with green space patch fragmentation. The phylogenetic diversity of both cultivated and spontaneous plants were weakly associated with green space structure, which was strongly driven by land use. Old UFUs and those with larger populations had more green space patches overall, although they tended to be small and fragmented. Green space patch density was found to increase as the UFU age increased. From the viewpoint of knowledge transfer, understanding the effects and drivers of landscape patterns of urban green spaces could inform the development of improved policies and management of urban green space areas.
- Open Access
- 9 July 2020
- Published by MDPI AG in Remote Sensing
- Vol. 12 (14) , 2201
An Integrated Approach to Study Spatial Patterns and Drivers of Land Cover Within Urban Functional Units: A Multi-City Comparative Study in China
Understanding the factors that drive green space composition and richness in heterogeneous urban landscapes is critical for maintaining important ecosystem services and biodiversity. Few studies have been conducted on urban greening and plant diversity at the urban functional unit (UFU) level, although a handful of studies have explored the drivers of greening percentage and its relationships with plant richness in tropical cities. In this study, we conducted field surveys, compiled census and remote sensing data, and performed spatial analyses to investigate the interrelationship between greening percentages, plant diversity, and the socioeconomic variables of different primary and secondary UFUs in the cities of Beijing, Zhanjiang, and Haikou in China. We found that these relationships did not differ significantly between primary and secondary UFUs, and that Parks represented the largest areas of forested land, grassland, and water bodies across all three cities. Moreover, the greening percentages of all UFUs across these three cities were positively correlated with both socioeconomic variables and plant species richness. These relationships can be utilized to guide future green space planning within urban ecosystems.
- 9 July 2020
- Published by MDPI AG in Remote Sensing
- Vol. 12 (14) , 2201
Open AccessAn Integrated Approach to Study Spatial Patterns and Drivers of Land Cover Within Urban Functional Units: A Multi-City Comparative Study in China
Understanding the factors that drive green space composition and richness in heterogeneous urban landscapes is critical for maintaining important ecosystem services and biodiversity. Few studies have been conducted on urban greening and plant diversity at the urban functional unit (UFU) level, although a handful of studies have explored the drivers of greening percentage and its relationships with plant richness in tropical cities. In this study, we conducted field surveys, compiled census and remote sensing data, and performed spatial analyses to investigate the interrelationship between greening percentages, plant diversity, and the socioeconomic variables of different primary and secondary UFUs in the cities of Beijing, Zhanjiang, and Haikou in China. We found that these relationships did not differ significantly between primary and secondary UFUs, and that Parks represented the largest areas of forested land, grassland, and water bodies across all three cities. Moreover, the greening percentages of all UFUs across these three cities were positively correlated with both socioeconomic variables and plant species richness. These relationships can be utilized to guide future green space planning within urban ecosystems.
- Open Access
- 25 June 2020
- Published by Frontiers Media SA in Frontiers in Plant Science
- Vol. 11, 941
Multiple MYB Activators and Repressors Collaboratively Regulate the Juvenile Red Fading in Leaves of Sweetpotato
Juvenile red fading describes the phenomenon in plants whereby red young leaves gradually turn green as they mature. While this phenomenon is commonly observed, the underlying molecular mechanism is still obscure as the classic model plants do not exhibit this process. Here, the molecular mechanism for the loss of anthocyanins during juvenile red fading were explored in the sweetpotato (Ipomoea batatas L.) cultivar “Chuanshan Zi”. The MYB-bHLH-WDR (MBW) regulatory complexes for anthocyanins were examined with five stages of leaf development from C1 to C5. Alternating accumulation of anthocyanins and chlorophylls caused the leaf color change. Five anthocyanin components were identified by ultra performance liquid chromatography/tandem mass spectrometry (UPLC-MS/MS), and their contents were highest at stage C2. Transcriptomic analysis showed massive gene expression alteration during leaf development. The anthocyanin structural genes expressed in sweetpotato leaves were screened and found to be highly comparable with those identified in morning glories. The screened anthocyanin regulatory genes included one bHLH (IbbHLH2), one WDR (IbWDR1), three MYB activators (IbMYB1, IbMYB2, and IbMYB3), and five MYB repressors (IbMYB27, IbMYBx, IbMYB4a, IbMYB4b, and IbMYB4c). The expression trends of MYBs were key to the red fading process: the activators were highly expressed in early red leaves and were all accompanied by simultaneously expressed MYB repressors, which may act to prevent excessive accumulation of anthocyanins. The only antagonistic repressor, IbMYB4b, was highly expressed in green leaves, and may be critical for declined anthocyanin content at later stages. Further functional verification of the above transcription factors were conducted by promoter activation tests. These tests showed that the MBW complexes of IbMYB1/IbMYB2/IbMYB3-IbbHLH2-IbWDR1 not only activated promoters of anthocyanin structural genes IbCHS-D and IbDFR-B, but also promoters for IbbHLH2 and IbMYB27, indicating both hierarchical and feedback regulations. This study outlines the elaborate regulatory network of MBW complexes involving multiple MYBs which allow for the timely accumulation of anthocyanins in sweetpotato leaves. These results may also provide clues for similar studies of juvenile red fading in other plant species.
- 25 June 2020
- Published by Frontiers Media SA in Frontiers in Plant Science
- Vol. 11, 941
Open AccessMultiple MYB Activators and Repressors Collaboratively Regulate the Juvenile Red Fading in Leaves of Sweetpotato
Juvenile red fading describes the phenomenon in plants whereby red young leaves gradually turn green as they mature. While this phenomenon is commonly observed, the underlying molecular mechanism is still obscure as the classic model plants do not exhibit this process. Here, the molecular mechanism for the loss of anthocyanins during juvenile red fading were explored in the sweetpotato (Ipomoea batatas L.) cultivar “Chuanshan Zi”. The MYB-bHLH-WDR (MBW) regulatory complexes for anthocyanins were examined with five stages of leaf development from C1 to C5. Alternating accumulation of anthocyanins and chlorophylls caused the leaf color change. Five anthocyanin components were identified by ultra performance liquid chromatography/tandem mass spectrometry (UPLC-MS/MS), and their contents were highest at stage C2. Transcriptomic analysis showed massive gene expression alteration during leaf development. The anthocyanin structural genes expressed in sweetpotato leaves were screened and found to be highly comparable with those identified in morning glories. The screened anthocyanin regulatory genes included one bHLH (IbbHLH2), one WDR (IbWDR1), three MYB activators (IbMYB1, IbMYB2, and IbMYB3), and five MYB repressors (IbMYB27, IbMYBx, IbMYB4a, IbMYB4b, and IbMYB4c). The expression trends of MYBs were key to the red fading process: the activators were highly expressed in early red leaves and were all accompanied by simultaneously expressed MYB repressors, which may act to prevent excessive accumulation of anthocyanins. The only antagonistic repressor, IbMYB4b, was highly expressed in green leaves, and may be critical for declined anthocyanin content at later stages. Further functional verification of the above transcription factors were conducted by promoter activation tests. These tests showed that the MBW complexes of IbMYB1/IbMYB2/IbMYB3-IbbHLH2-IbWDR1 not only activated promoters of anthocyanin structural genes IbCHS-D and IbDFR-B, but also promoters for IbbHLH2 and IbMYB27, indicating both hierarchical and feedback regulations. This study outlines the elaborate regulatory network of MBW complexes involving multiple MYBs which allow for the timely accumulation of anthocyanins in sweetpotato leaves. These results may also provide clues for similar studies of juvenile red fading in other plant species.