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Hydrastis Canadensis L.) Phytochemistry, Trade, and Habitat

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Hydrastis Canadensis L.) Phytochemistry, Trade, and Habitat The Pennsylvania State University The Graduate School GOLDENSEAL (HYDRASTIS CANADENSIS L.) PHYTOCHEMISTRY, TRADE, AND HABITAT: IMPLICATIONS FOR CONSERVATION AND FOREST BASED CULTIVATION A Dissertation in Forest Resources by Grady Henry Zuiderveen © 2019 Grady Henry Zuiderveen Submitted in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy August 2019 The dissertation of Grady Zuiderveen was reviewed and approved* by the following: Eric Burkhart Plant Science Program Director, Shaver’s Creek Environmental Center Dissertation Co-Advisor Co-Chair of Committee Michael Jacobson Professor of Forest Resources Dissertation Co-Advisor Co-Chair of Committee Margot Kaye Associate Professor of Forest Ecology Joshua Lambert Associate Professor of Food Science Co-Director, Center for Plant and Mushroom Foods for Health Doug Miller Research Professor of Geography Director, Center for Environmental Informatics Michael Messina Head and Professor, Department of Ecosystem Science and Management *Signatures are on file in the Graduate School ii Abstract Goldenseal (Hydrastis canadensis L.) is a well-known medicinal, forest plant native to eastern North America. The roots and rhizomes of this species are used primarily as an antimicrobial and for the treatment of inflammation and intestinal ailments. Due to conservation concerns over the long-term viability of wild populations, the species has been listed in Appendix 2 of the Convention on International Trade of Endangered Species of Wild Fauna and Flora (CITES), an international trade treaty surrounding wild species. Concurrently, there is demand in the herbal industry for high quality, sustainably sourced goldenseal products. This study examined phytochemistry of wild harvested rhizomes and shoots; behaviors and perspectives surrounding goldenseal trade in the eastern United States; and habitat suitability and indicators in Pennsylvania. This interdisciplinary set of studies was undertaken to better understand and enhance wild conservation efforts, and to develop information that could be used to inform the adoption of cultivation, forest farming, and in situ wild stewardship. Phytochemistry studies were undertaken to investigate the occurrence of three isoquinoline alkaloids—berberine, hydrastine, and canadine—currently recognized as three major bioactive constituents in goldenseal. High performance liquid chromatography (HPLC) was used to look at the influence of development stage, time-of-day, habitat conditions, and drying temperature on variations in alkaloid concentrations in goldenseal rhizomes and shoots. Results indicate that harvest and post-harvest conditions and practices can greatly influence the total alkaloid concentration in the plant. Alkaloid concentration was found to peak in both plant shoots and in rhizomes at flowering stage and in the rhizomes in the dormancy stage. Hydrastine and canadine levels were found to be greatest in shoots in the afternoon. Habitat factors such as forest overstory iii canopy cover, plant density, and soil cation exchange capacity (CEC) were all found to influence alkaloid levels in goldenseal. Drying temperatures (26.7°C – 54.4°C) were found to influence canadine, which decreased as temperature increased. To better understand goldenseal trade behaviors and conservation perspectives, surveys were used to gather information around the management, trade, and conservation practices in Pennsylvania, and across the species natural range. Results indicate that trade is more significant and widespread in goldenseal’s core range (Ohio, Indiana, Kentucky, and West Virginia), and that this increased trade activity is also linked to perceived greater availability of goldenseal as a wild species and less concern by buyers over conservation. Further, there is a lack of state involvement in goldenseal trade tracking and management when compared with American ginseng, which is another CITES listed species. Results indicate that the state-to-state variation in harvest may require greater attention at the state, rather than federal, level to improve a regionally-focused approach to goldenseal conservation. Lastly, indicator species analysis and habitat suitability modeling were used to identify goldenseal habitat in Pennsylvania – a state at the northern edge of its range. The results identified bedrock type and the average winter temperature as important factors in determining suitable habitat. The most prevalent overstory tree associates with goldenseal populations were tulip-poplar (Liriodendron tulipifera) and sugar maple (Acer saccharum); common understory associates included spicebush (Lindera benzoin), Jack-in-the-pulpit (Arisaema triphyllum), and rattlesnake fern (Botrypus virginianus). Taken together, these results identified rich, mesic, woodland sites in the south and south-central part of the state as being suitable for goldenseal, and by prioritizing these areas for protection, management, and cultivation (i.e., forest farming), greater levels of conservation success could be achieved. iv TABLE OF CONTENTS List of Figures…………………………………………………………………………………………………………………… viii List of Tables……………………………………………………………………………………………………………………. xi Acknowledgments…………………………………………………………………………………………………………… xiii Chapter 1 Introduction……………………………………………………………………………………………………. 1 Botany………………….……………………………………………………………………………………………… 1 Medicinal Use..……………………………………………………………………………………………………. 2 Quality Control……………………………………………………………………………………………………. 4 Non-Timber Forest Products.………………………………………………………………………………. 5 Trade Status….……………………………………………………………………………………………………… 6 Conservation Concerns………………………………………………………………………………………… 7 In Situ Stewardship and Forest-Based Cultivation………………..………………………………. 8 Dissertation Objectives………………………………………………………………………………………… 11 Literature Cited……………………………………………………………………………………………………. 12 Chapter 2 Influence of post-harvest drying temperatures on alkaloid levels in goldenseal (Hydrastis canadensis L.) ……………………………….……………......…………… 23 Abstract…………………………………………………………………………………………………………….... 23 Introduction…………………………………………………………………………………………………………. 24 Materials and Methods………………………………………………………………………………………… 26 Plant Material…………………………………………………………………………………………… 26 Processing Conditions.……………………………………………………………………………… 26 Chemical Calibration Standards………………………………………………………………… 27 HPLC Analysis…………………………………………………………………………………………… 27 Statistical Analysis……………………………………………………………………………………. 28 Results and Discussion…………………………………………………………………………………………. 28 Literature Cited……………………………………………………………………………………………………. 30 Chapter 3 Effects of phenological stage, time-of-day, and forest site conditions on alkaloid content in goldenseal (Hydrastis canadensis L.) rhizomes and shoots……. 39 Abstract…………………………………………………………………………………………………………….... 39 Introduction…………………………………………………………………………………………………………. 40 Materials and Methods………………………………………………………………………………………… 42 Plant Material…………………………………………………………………………………………… 42 Post-Harvest Processing…………………………………………………………………………… 44 Chemical Calibration Standards………………………………………………………………… 44 Sample Preparation………………………………………………………………………………….. 45 HPLC Analysis…………………………………………………………………………………………… 45 Soil Analysis……………………………………………………………………………………………… 45 Canopy Cover…………………………………………………………………………………………… 46 Plant Density.…………………………………………………………………………………………… 46 v Statistical Analysis……………………………………………………………………………………. 46 Results…………………………………………………………………………………………………………………. 47 Changes in Alkaloids with Phenological Stage…………………………………………… 47 Changes in Alkaloids with Time of Day……………………………………………………… 48 Changes in Alkaloids with Site Conditions………….…………………………………….. 48 Changes in Alkaloids and Dry Weight with Plant Morphology…………………… 48 Changes in Dry Weight with Phenological Stage………………………………………. 49 Discussion……………………………………………………………………………………………………………. 49 Implications for Wild Harvesting and Forest-Based Cultivation (i.e. Forest Farming) ………………………………………………………………………………… 49 Phenology and Secondary Metabolites in Plants………………………………………. 51 Site Conditions and Secondary Metabolites in Plants….……………………………. 52 Time of Day and Secondary Metabolites in Plants……...……………………………. 54 Conclusion…………………………………………………………………………………………………………… 54 Literature Cited……………………………………………………………………………………………………. 55 Chapter 4 Goldenseal (Hydrastis canadensis L.) trade practices and stakeholder perceptions in the United States: implications for management and conservation………………………………………………………………………………………………………… 72 Abstract…………………………………………………………………………………………………………….... 72 Introduction…………………………………………………………………………………………………………. 73 Materials and Methods....………………………………….………………………………………………… 77 Sampling Frame…………….…………………………….…………………………………………… 77 Survey Implementation...…………………………….…………………………………………… 78 Survey Analysis………….….…………………………….…………………………………………… 79 Results and Discussion…………………………………………………………………………………………. 80 Demand and Source for Goldenseal.………………………………………………………… 80 The Importance of Geography on Buying and Trading Practices………………. 83 Perspectives on Conservation and Regulation.…………………………………………. 84 State Efforts to Track, Conserve, and Regulate………………………………………… 87 Conclusion…………………………………………………………………………………………………………… 88 Literature Cited……………………………………………………………………………………………………. 89 Chapter 5 Associated habitat and suitability modeling of goldenseal (Hydrastis canadensis L.) in Pennsylvania: explaining and predicting species distribution in a northern edge of range state..……….……………………………………………………………… 114 Abstract…………………………………………………………………………………………………………….... 114 Introduction………………………………………………………………………………………………………….
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