Modelling the impact of future climate and land use change on vegetation patterns, plant diversity and provisioning ecosystem services in West Africa

Global climate change and land use change will not only alter entire ecosystems and biodiversity patterns, but also the supply of ecosystem services. A better understanding of the consequences is particularly needed in under-investigated regions, such as West Africa. The projected environmental changes suggest negative impacts on nature, thus representing a threat to the human well-being. However, many effects caused by climate and land use change are poorly understood so far. Thus, the main objective of this thesis was to investigate the impact of climate and land use change on vegetation patterns, plant diversity and important provisioning ecosystem services in West Africa. The three different aspects are separately explored and build the chapters of this thesis. The findings help to improve our understanding of the effects of environmental change on ecosystems and human well-being. In the first study, the main objectives were to model trends and the extent of future biome shifts in West Africa that may occur by 2050. Also, I modelled a trend in West African tree cover change, while accounting for human impact. Additionally, uncertainty in future climate projections was evaluated to identify regions with reliable trends and regions where the impacts remain uncertain. The potential future spatial distributions of desert, grassland, savanna, deciduous and evergreen forest were modelled in West Africa, using six bioclimatic models. Future tree cover change was analysed with generalized additive models (GAMs). I used climate data from 17 general circulation models (GCMs) and included human population density and fire intensity to model tree cover. Consensus projections were derived via weighted averages to: 1) reduce inter-model variability, and 2) describe trends extracted from different GCM projections. The strongest predicted effect of climate change was on desert and grasslands, where the bioclimatic envelope of grassland is projected to expand into the Sahara desert by an area of 2 million km2. While savannas are predicted to contract in the south (by 54 ± 22 × 104 km2), deciduous and evergreen forest biomes are expected to expand (64 ± 13 × 104 km2 and 77 ± 26 × 104 km2). However, uncertainty due to different GCMs was particularly high for the grassland and the evergreen forest biome shift. Increasing tree cover (1–10%) was projected for large parts of Benin, Burkina Faso, Côte d’Ivoire, Ghana and Togo, but a decrease was projected for coastal areas (1–20%). Furthermore, human impact negatively affected tree cover and partly changed the direction of the projected climate-driven tendency from increase to decrease. Considering climate change alone, the model results of potential vegetation (biomes) showed a ‘greening’ trend by 2050. However, the modelled effects of human impact suggest future forest degradation. Thus, it is essential to consider both climate change and human impact in order to generate realistic future projections on woody cover. The second study focused on the impact and the interplay of future (2050) climate and land use change on the plant diversity of the West African country Burkina Faso. Synergistic forecasts for this country are lacking to date. Burkina Faso covers a broad bioclimatic gradient which causes a similar gradient in plant diversity. Thus, the impact of climate and land use change can be investigated in regions with different levels of species richness. The LandSHIFT model from the Centre of Environmental System research CESR (Kassel, Germany) was adapted for this study to derive novel regional, spatially explicit future (2050) land use simulations for Burkina Faso. Additionally, the simulations include different assumptions on the technological developments in the agricultural sector. Oneclass support vector machines (SVMs), a machine learning method, were performed with these land use simulations together with current and future (2050) climate projections at a 0.1° resolution (cell: ~ 10 × 10 km). The modelling results showed that the flora of Burkina Faso will be primarily negatively impacted by future climate and land use changes. The species richness will be significantly reduced by 2050 (P < 0.001, paired Wilcoxon signed-rank test). However, contrasting latitudinal patterns were found. Although climate change is predicted to cause species loss in the more humid regions in Southern Burkina Faso (~ 200 species per cell), the model projects an increase of species richness in the Sahel. However, land use change is expected to suppress this increase to the current species diversity level, depending on the technological developments. Climate change is a more important threat to the plant diversity than land use change under the assumption of technological stagnation in the agricultural sector. Overall, the study highlights the impact and interplay of future climate and land use change on plant diversity along a broad bioclimatic gradient in West Africa.Furthermore, the results suggest that plant diversity in dry and humid regions of the tropics might generally respond differently to climate and land use change. This pattern has not been detected by global studies so far. Several of the plant species in West Africa significantly contribute to the livelihoods of the population. The plants provide so-called non-timber forest products (NTFPs), which are important provisioning ecosystem services. However, these services are also threatened by environmental change. Thus, the third study aimed at developing a novel approach to assess the impacts of climate and land use change on the economic benefits derived from NTFPs. This project was carried out in cooperation with Katja Heubach (BiK-F) who provided data on household economics. These data include 60 interviews that were conducted in Northern Benin on annual quantities and revenues of collected NTFPs from the three most important savanna tree species: Adansonia digitata, Parkia biglobosa and Vitellaria paradoxa. The current market prices of the NTFPs were derived from respective local markets. To assess current and future (2050) occurrence probabilities of the three species, I calibrated niche-based models with climate data (from Miroc3.2medres) and land use data (LandSHIFT) at a 0.1° resolution (cell: ~ 10 × 10 km). Land use simulations were taken from the previous study on plant diversity. Three different niche-based models were used: 1) generalized additive models (regression method), 2) generalized boosting models (machine learning method), and 3) flexible discriminant analysis (classification method). The three model simulations were averaged (ensemble forecasting) to increase the robustness of the predictions. To assess future economic gains and losses, respectively, the modelled species’ occurrence probabilities were linked with the spatially assigned monetary values. Highest current annual benefits are obtained from V. paradoxa (54,111 ± 28,126 US$/cell), followed by P. biglobosa (32,246 ± 16,526 US$/cell) and A. digitata (9,514 ± 6,243 US$/cell). However, in the prediction large areas will lose up to 50$/Gridzelle), gefolgt von Parkia biglobosa (32.246 ± 16.526 US$/Gridzelle) und Adansonia digitata (9.514 ± 6.243 US$/Gridzelle). Allerdings zeigen die Zukunftsprojektionen für das Jahr 2050, dass große Flächen in Nordbenin bis zu 50% dieses ökonomischen Wertes verlieren könnten. Am stärksten betroffen sind dabei die Arten Vitellaria paradoxa und Parkia biglobosa, die derzeit den höchsten Nutzwert haben. Adansonia digitata wird weniger stark aber ebenfalls negativ beeinflusst werden in weiten Teilen von Nordbenin. Hier zeigen sich jedoch regional ausgeprägte Unterschiede. Vor allem im Westen und im Osten des Untersuchungsgebietes können die Umweltveränderungen zu erhöhten Auftrittswahrscheinlichkeiten und damit höheren Erlösen im Jahr 2050 führen. Insgesamt zeigen die Ergebnisse, dass Adaptationsmaßnahmen erforderlich sind um alternative Einkommensquellen zu erschließen. Dies gilt insbesondere für Frauen, die für das Sammeln der NTFPs verantwortlich sind. Die Ergebnisse liefern Politikern eine Orientierungsmöglichkeit um verschiedene Landnutzungsoptionen ökonomisch zu vergleichen und etwaige Anpassungen von rezenten

Securin Is Not Required for Chromosomal Stability in Human Cells

Abnormalities of chromosome number are frequently observed in cancers. The mechanisms regulating chromosome segregation in human cells are therefore of great interest. Recently it has been reported that human cells without an hSecurin gene lose chromosomes at a high frequency. Here we show that, after hSecurin knockout through homologous recombination, chromosome losses are only a short, transient effect. After a few passages hSecurin(−/−) cells became chromosomally stable and executed mitoses normally. This was unexpected, as the securin loss resulted in a persisting reduction of the sister-separating protease separase and inefficient cleavage of the cohesin subunit Scc1. Our data demonstrate that securin is dispensable for chromosomal stability in human cells. We propose that human cells possess efficient mechanisms to compensate for the loss of genes involved in chromosome segregation

The AAA-ATPase p97 in mitosis and fertilization

Late mitotic events are chiefly controlled by proteolysis of key regulatory proteins via the ubiquitin-proteasome pathway. In this pathway ubiquitin ligases modify substrates by attachment of ubiquitin (“ubiquitylation”), which usually results in their subsequent degradation by the 26S proteasome. The crucial ubiquitin ligase involved in late mitosis is the anaphase-promoting complex or cyclosome (APC/C). Among the many substrates of the APC/C is the anaphase inhibitor securin, whose destruction leads to activation of separase, which in turn triggers sister chromatid separation by proteolytic cleavage of cohesin. The APC/C also targets cyclin B1, an activating subunit of Cdk1 kinase, whose inactivation is a prerequisite for mitotic exit. The unstable APC/C substrates are often found in association with stable partner proteins. How single subunits of multi-protein complexes are selectively extracted and eventually degraded is largely unknown, but there is increasing evidence that additional factors assist to extract ubiquitin-carrying subunits from stable binding partners. One such factor is vertebrate p97 (Cdc48 in yeast), an abundant and highly conserved member of the AAA-ATPase family. It is involved in such diverse processes as transcriptional regulation, membrane fusion, and ER-associated protein degradation (ERAD). The unifying scheme in these seemingly unrelated functions is that p97 is able to “extract” preferentially ubiquitylated proteins from their environment. Roles of p97 in mitosis have recently emerged: p97 was reported to be required for spindle disassembly and for nuclear envelope reformation during mitotic exit in Xenopus. Furthermore, a genetic interaction between p97, separase and securin, as well as a requirement of p97 for separase stability, were discovered in fission yeast. Given these hints and the importance of ubiquitylation in both mitosis and p97 pathways, this study intended to elucidate additional mitotic roles of p97 in vertebrates. Towards this end, tools to interfere with p97 function in Xenopus egg extracts were developed. These included immunodepletion of the p97 adaptors Npl4, Ufd1 and p47 and addition of recombinant dominant-negative p97-mutants. ERAD, which could be established here for the first time in Xenopus egg extracts, was greatly impaired in the absence of p97 function. However, many aspects of mitosis were found to be unaffected. Importantly, p97’s proposed role in spindle disassembly was clearly falsified within this thesis. Furthermore, p97 was shown to be dispensable for activity and stability of vertebrate separase. Disassembly of the mitotic checkpoint complex, which prevents premature APC/C activation by sequestering its activator Cdc20, did also not require functional p97 despite its dependence on ubiquitylation of Cdc20. However, a novel function of p97 at fertilization was discovered. p97 was found to interact with nucleoplasmin, a histone-binding chaperone that catalyzes the exchange of sperm-specific basic proteins (SBPs) to histones. Indeed, interference with p97 function delayed sperm decondensation in Xenopus egg extracts, thereby confirming a novel role of this AAA-ATPase in sperm chromatin remodelling. In another project the role of securin in human cells was investigated. Human cells lacking securin had been reported to suffer from massive chromosome missegregation, which was in sharp contrast to the mild phenotype of securin knockout mice. In collaboration with the group of M. Speicher it could be demonstrated that chromosome losses in securin-/- cells are transient and give way to a stable segregation pattern after just a few passages. This was despite persisting biochemical defects such as reduced level and activity of separase. These data demonstrate that securin is dispensable for chromosomal stability in human cells

CUL-2^LRR-1 and UBXN-3 drive replisome disassembly during DNA replication termination and mitosis

Replisome disassembly is the final step of DNA replication in eukaryotes, involving the ubiquitylation and CDC48-dependent dissolution of the CMG helicase (CDC45-MCM-GINS). Using Caenorhabditis elegans early embryos and Xenopus laevis egg extracts, we show that the E3 ligase CUL-2(LRR-1) associates with the replisome and drives ubiquitylation and disassembly of CMG, together with the CDC-48 cofactors UFD-1 and NPL-4. Removal of CMG from chromatin in frog egg extracts requires CUL2 neddylation, and our data identify chromatin recruitment of CUL2(LRR1) as a key regulated step during DNA replication termination. Interestingly, however, CMG persists on chromatin until prophase in worms that lack CUL-2(LRR-1), but is then removed by a mitotic pathway that requires the CDC-48 cofactor UBXN-3, orthologous to the human tumour suppressor FAF1. Partial inactivation of lrr-1 and ubxn-3 leads to synthetic lethality, suggesting future approaches by which a deeper understanding of CMG disassembly in metazoa could be exploited therapeutically

Genetic Signatures of a Demographic Collapse in a Large-Bodied Forest Dwelling Primate (Mandrillus Leucophaeus)

It is difficult to predict how current climate change will affect wildlife species adapted to a tropical rainforest environment. Understanding how population dynamics fluctuated in such species throughout periods of past climatic change can provide insight into this issue. The drill (Mandrillus leucophaeus) is a large-bodied rainforest adapted mammal found in West Central Africa. In the middle of this endangered monkey’s geographic range is Lake Barombi Mbo, which has a well-documented palynological record of environmental change that dates to the Late Pleistocene. We used a Bayesian coalescent-based framework to analyze 2,076 base pairs of mitochondrial DNA across wild drill populations to infer past changes in female effective population size since the Late Pleistocene. Our results suggest that the drill underwent a nearly 15-fold demographic collapse in female effective population size that was most prominent during the Mid Holocene (approximately 3-5 Ka). This time period coincides with a period of increased dryness and seasonality across Africa and a dramatic reduction in forest coverage at Lake Barombi Mbo.We believe that these changes in climate and forest coverage were the driving forces behind the drill population decline. Furthermore, the warm temperatures and increased aridity of the Mid Holocene are potentially analogous to current and future conditions faced by many tropical rainforest communities. In order to prevent future declines in population size in rainforest-adapted species such as the drill, large tracts of forest should be protected to both preserve habitat and prevent forest loss through aridification. 55

Cdc48 and Cofactors Npl4-Ufd1 Are Important for G1 Progression during Heat Stress by Maintaining Cell Wall Integrity in Saccharomyces cerevisiae

The ubiquitin-selective chaperone Cdc48, a member of the AAA (ATPase Associated with various cellular Activities) ATPase superfamily, is involved in many processes, including endoplasmic reticulum-associated degradation (ERAD), ubiquitin- and proteasome-mediated protein degradation, and mitosis. Although Cdc48 was originally isolated as a cell cycle mutant in the budding yeast Saccharomyces cerevisiae, its cell cycle functions have not been well appreciated. We found that temperature-sensitive cdc48-3 mutant is largely arrested at mitosis at 37°C, whereas the mutant is also delayed in G1 progression at 38.5°C. Reporter assays show that the promoter activity of G1 cyclin CLN1, but not CLN2, is reduced in cdc48-3 at 38.5°C. The cofactor npl4-1 and ufd1-2 mutants also exhibit G1 delay and reduced CLN1 promoter activity at 38.5°C, suggesting that Npl4-Ufd1 complex mediates the function of Cdc48 at G1. The G1 delay of cdc48-3 at 38.5°C is a consequence of cell wall defect that over-activates Mpk1, a MAPK family member important for cell wall integrity in response to stress conditions including heat shock. cdc48-3 is hypersensitive to cell wall perturbing agents and is synthetic-sick with mutations in the cell wall integrity signaling pathway. Our results suggest that the cell wall defect in cdc48-3 is exacerbated by heat shock, which sustains Mpk1 activity to block G1 progression. Thus, Cdc48-Npl4-Ufd1 is important for the maintenance of cell wall integrity in order for normal cell growth and division