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Journal of Resources and Ecology >

2022 , Vol. 13 >Issue 5: 925 - 935

DOI: https://doi.org/10.5814/j.issn.1674-764x.2022.05.016

Assessing Impact of Restoration on Livelihood

A Review on the Supply-Demand Relationship and Spatial Flows of Ecosystem Services

  • HUANG Mengdong , 1, 2 ,
  • XIAO Yu , 1, 2, * ,
  • XU Jie 3 ,
  • LIU Jingya 1, 2 ,
  • WANG Yangyang 1, 2 ,
  • GAN Shuang 1, 2 ,
  • LV Shixuan 4 ,
  • XIE Gaodi 1, 2
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  • 1. Institute of Geographic Sciences and Natural Resources Research, Chinese Academy of Science, Beijing 100101, China
  • 2. University of Chinese Academy of Sciences, Beijing 100049, China
  • 3. School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083, China
  • 4. College of Geography and Environment, Shandong Normal University, Jinan 250358, China
*XIAO Yu, E-mail:

HUANG Mengdong, E-mail:

Received date: 2021-11-14

  Accepted date: 2022-03-02

  Online published: 2022-07-15

Supported by

The National Natural Science Foundation of China(41971272)

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Abstract

Research on spatial flow as it relates to the relationship between the supply and demand of ecosystem services supports a significant connection between the supply of ecosystem services and human well-being. Understanding the entire process of the production and flow, as well as the use of ecosystem services, accurately assessing the balance of supply and demand of ecosystem services, and establishing a two-way feedback relationship between supply and demand are vital for the scientific management of the ecosystem and ensuring the sustainable development of regional resources. Based on a large number of relevant publications, this paper comprehensively summarizes the concepts and assessment methods of ecosystem service supply and demand from the perspective of ecosystem service supply and demand, and discusses the impacts of land use and climate change on the temporal and spatial changes of ecosystem services under the background of global change. Then, an analysis of the research progress in the ecosystem services spatial flow indicated that there are still deficiencies in the quantification of cultural services, the dynamics of ecosystem service flow and the driving mechanism of ecosystem services. We also propose that clarifying the driving mechanism and transfer process of ecosystem services, and realizing the mutual conversion between different spatial-temporal scales of ecosystem services, is an important approach for improving the application of ecosystem services research in practice in the future.

Key words： ecosystem services; supply-demand relationship; spatial flow; value assessment; driving factors

Cite this article

HUANG Mengdong , XIAO Yu , XU Jie , LIU Jingya , WANG Yangyang , GAN Shuang , LV Shixuan , XIE Gaodi . A Review on the Supply-Demand Relationship and Spatial Flows of Ecosystem Services[J]. Journal of Resources and Ecology, 2022 , 13(5) : 925 -935 . DOI: 10.5814/j.issn.1674-764x.2022.05.016

1 Introduction

Since the 1990s, with the efforts from Daily (1997) and Costanza et al. (1997), ecosystem services have gradually developed from conceptual research to a new stage of systematic and comprehensive research. Many scholars have carried out a large number of value evaluation studies on different types of ecosystem services from various spatiotemporal scales based on ecology and economics (Xie et al., 2001; Sutton and Costanza, 2002; Shi et al., 2012). In 2005, the Millennium Ecosystem Assessment defined ecosystem services as the benefits that humans obtain from ecosystems, and emphasized the importance of human well-being in ecosystem services research (MEA, 2005). Therefore, human demand has been increasingly taken into account in the research on ecosystem services and has also mainly focused on the two main contents of whether the needs of human society were met and whether the balance of the supply and demand of ecosystem services was sustainable (Yahdjian et al., 2015; Chen et al., 2019; Wu et al., 2019; Zhao et al., 2019; Zhai et al., 2020). However, with the global change, ecosystem services are constantly changing due to the impacts from natural and human factors. The supply-demand relationship of ecosystem services is therefore affected, which has an adverse effect on the long-term and stable implementation of policies such as ecosystem services and natural resource management (Metzger and Schröter, 2006; Weiskopf et al., 2020). Although many previous studies have explored the responses of ecosystem services to land use and climate change, the reported correlations between ecosystem natural processes and human activities are complicated (Long et al., 2014; Scholes, 2016; Runting et al., 2017; Paudyal et al., 2019; Rimal et al., 2019), and the factors driving ecosystem service supply and demand and their influencing mechanisms are still unclear. In these regards, further studies are still necessary.
In addition, some scholars have revealed the characteristics of spatial mismatches between the supply and demand of ecosystem services, which means the supply area and supply capacity of services do not overlap with the demand area and degree of demand within the same spatial limit (Burkhard et al., 2012; Nedkov and Burkhard, 2012). With ecosystem service flow, the supply and demand of services are effectively connected, ensuring the entire process of ecosystems from generation to utilization. By clarifying the specific flow paths of ecosystem services, it has been possible to analyze the dynamic balance of ecosystem service supply and demand, to identify the key locations that are critical for the delivery of ecosystem services, to provide a scientific basis for formulating reasonable ecosystem management policies and ecological compensation plans, and to realize the sustainable development of regional ecological security. However, studies on the ecosystem services flow are still in the preliminary exploration stage, and the methods of path simulation and quantitative evaluation are not yet systematic. Despite these limitations, a few scholars have made attempts to simulate and quantify the spatial flows of some ecosystem services (Vallecillo et al., 2019; Xu et al., 2019; Shi et al., 2020), which have laid the foundation for the dynamic research of ecosystem services.
According to the ecosystem service supply and demand, this article summarizes the evaluation methods and driving factors of the ecosystem service supply and demand, and the correlations between them. In addition, we also generalize the concept, attributes and methods of ecosystem service flow, discuss the difficulties and problems in the current stage, and also point out the future research directions of ecosystem services.

2 Relationship between the supply and demand of ecosystem services

2.1 Ecosystem service supply and demand

The concept of ecosystem service supply was originally related to ecological carrying capacity, which is the supply capacity of resources in a specific area, also known as potential supply. The potential supply is provided by the ecosystem in a sustainable manner for the long term. However, because the ecosystem needs some energy to maintain its own balance, the ecosystem cannot offer all of the resources that humans need. Only products or ecological processes that are actually consumed by humans can be considered to represent ecosystem service supply, also known as actual supply (Ma et al., 2017). Burkhard et al. (2012) believed that the ecosystem service supply refers to the availability of specific ecosystem products or services that could be actually used in a specific area and provided for a specific period of time. The place where ecosystem services are produced is the ecosystem service supply area (Fisher et al., 2009). It is composed of ecosystems, populations and other physical components (Syrbe and Walz, 2012), which also affect the supply capacity of ecosystem services to varying degrees.
As the connections between ecosystem services and human well-being become closer and closer, the ecosystem service supply research has been gradually integrated with human production life. Therefore, the demand of ecosystem services has become another main concentration for the scholars who major in ecosystem services. At present, no clear definition of the ecosystem services demand has attained universal support. From the perspective of the ultimate direction of ecosystem services, Burkhard et al. (2012) defined ecosystem service requirements as ecosystem services used by humans in a specific time and space, while Villamagna et al. (2013) defined and quantified ecosystem service demand from the perspective of service beneficiaries. She thought that ecosystem service demand was the quantity and quality of ecosystem services needed by humans. Based on the views of multiple scholars, the ecosystem service demand is closely related to human society. The ecosystem services that are effectively consumed or used by humans are called ecosystem service demand (Li et al., 2012; Bagstad et al., 2013; Liu et al., 2017). Corresponding to the ecosystem service supply, Ma et al. (2017) divided the definition of ecosystem service demand into two categories: actual demand and total demand. The former is the human demand for ecosystem services that could be met, while the latter is the unsatisfied demand. The areas where ecosystem services are used or consumed are called ecosystem service demand areas (Syrbe and Walz, 2012).
Synthesizing the concepts of ecosystem service supply and demand shows that ecosystem services are a complex process of coupling nature and humans, with significant attributes of timeliness, space and availability. The supply of ecosystem services is usually restricted by the spatial-temporal distribution and supply capacity of the natural processes of the ecosystem. Similarly, the demand for ecosystem services is more strongly affected by the degree of human dependence on resources, subjective preferences and consumption capacity. Therefore, ecosystem services also have significant spatiotemporally differentiated characteristics. In a given spatiotemporal area, the human demand for ecosystem services is not completely consistent with the supply capacity. Thus, it is necessary to adjust the supply structure of ecosystem services in different regions according to human needs and preferences in order to achieve the long-term balanced development of ecosystem services.

2.2 Evaluation methods for the supply and demand of ecosystem services

Research on the supply of ecosystem services started several decades ago. Material quantity simulations and value assessments are commonly used methods for evaluating the supply of ecosystem services. Based on the perspective of the ecosystem, the former reveals the mechanisms of ecological processes and simulates the quantities of ecosystem services by using models. The latter mostly simulates the value of ecosystem services through methods such as field surveys and market surveys, and it converts the quantities into values using models. The study areas of the research on the supply of ecosystem services abroad have mostly been natural geographic units, such as river basins or mountains, while those of the domestic research are usually administrative divisions. In addition, Koellner et al. (2019) used open and regional geographic thinking as a research method of system boundaries, instead of the traditional geographic boundaries of ecosystem service assessment, and explored the impacts on services of the external factors involved in the flow of ecosystems between regions.
In recent years, with the continuous in-depth study of ecosystem service value evaluation, many studies have begun to pay attention to the role of ecosystem service demand in ecosystem management. This research analyzes the characteristics of the spatiotemporal distribution of ecosystem service demand according to the distribution and the degree of stakeholders. Different scholars have explored different types of services. For example, Kroll et al. (2012) simulated food supply services by service consumption and the spatial distribution of beneficiaries; Nedkov and Burkhard (2012) simulated the demand for flood regulating services according to the demand levels in different kinds of land cover. However, since the demand indicators that were commonly used, such as population, GDP, land use degree, night lighting data, road network density, etc., did not completely represent the spatial heterogeneity of the ecosystem service demand, or the indicators themselves were difficult to quantify, it has been difficult to select suitable indices in such studies (Gu et al., 2018; Wang et al., 2018; Wang et al., 2019; Guo et al., 2020). Therefore, research on the demand for ecosystem services is still lacking.
With the continuous development of remote sensing and GIS, more and more studies are considering the spatial relationship between the supply and demand of ecosystem services. The quantification and spatial simulation of the supply and demand of ecosystem services has gradually become the main research method for determining the balance of the supply and demand of ecosystem services. At present, there are model simulation methods, value assessments, questionnaires and other methods for the comprehensive analysis of the balance between supply and demand of ecosystem services (Table 1). Among them, the model simulation method has mainly been used for research on the supply of ecosystem services and the balance between supply and demand, although this method relies on data with higher accuracy and needs complex calculation processes. However, taking the human needs as the research object, the participation method is applicable to the assessment of ecosystem service needs. The value evaluation method is more applicable in these two aspects, but it has been mostly applied to the types of ecosystem services with markets, and the results are greatly influenced by the subjective influences of experts. At the same time, the model simulation and value evaluation methods both ignore the characteristics of internal differences to varying degrees. In addition, due to human related factors, there are still relatively few studies on cultural services and the demand of ecosystem services.
Table 1 Comparisons of the main ecosystem service supply and demand evaluation methods
Evaluation types Category Method Advantages Limitations Ecosystem services types Sources
Ecosystem service
supply		 Simulation modeling InVEST Data is easy to obtain; results are visualized Simplify the algorithm, the results have errors; unable to obtain the information flow between some factors; strong dependence on data Provision services; regulation services; culture services Dong et al., 2018; Chen et al., 2019
CASA It can be applied widely; data is easy to obtain Due to the flaws in the data itself, the parameter correction effect is poor Provision services;
Regulation services		 Wei et al., 2018
RUSLE It can reveal the mechanisms of ecological processes; It can be applied widely Data dependence is strong and difficult to obtain Regulation services Castillo-Eguskitza et al., 2018
SWAT Data is easy to obtain Data accuracy requirements are high; there are many parameters and the parameter calibration is uncertain Regulation services Nedkov and Burkhard, 2012;
Yan et al., 2019
SoLVES The evaluation result is more accurate; It can be applied widely A large amount of field investigation and research are required to ensure the accuracy of the data; the value estimation ignores the differences within the study area Culture services Ma et al., 2018;
Zhao et al., 2019
Value assessment Travel cost It can indicate the preferences of the beneficiaries Only assesses part of the use value Culture services Li and Li, 2003;
Sun et al., 2017
Alternative cost Able to evaluate services that do not have market value through alternatives Only applicable to services with alternatives Regulation services Li and Zhou, 2016
Recovery
protection
expenditure		 Weak data dependency Can only evaluate the lowest value of the ecological environment Regulation services Morri et al., 2014
Ecosystem service demand Participate
survey		 Questionnaire survey It can be used in potential demand Subjectively affected by users; heavy workload Provision services; regulation services; culture services Wei et al., 2018; Quintas-Soriano et al., 2019




Ecosystem service supply and demand		 Simulation modeling ARIES High evaluation accuracy Mainly applicable to the United States; higher data requirements, high-resolution spatial data is required Provision services; regulation services; culture services Burkhard et al., 2012; Martinez-Lopez et al., 2019
Value assessment Value equivalent method Quantifies the value of different service functions;
avoids subjectivity		 Ignores intra-regional differences Provision services; regulation services; culture services Xie et al., 2015;
Gu et al., 2018;
Wang et al., 2018
Direct market value method Data is easy to obtain Only evaluates services that have market Provision services Castillo-Eguskitza
et al., 2018
Conditional Value Method It can be applied widely Based on a virtual market, with certain subjectivity and uncertainty Provision services; regulation services; culture services Burkhard et al., 2012; Castillo-Eguskitza
et al., 2018
Expert scoring Data is easy to obtain Subjectively influenced by experts Provision services; regulation services; culture services Ou et al., 2018;
Wu et al., 2019
The quantitative simulation of the supply and demand of ecosystem services requires diverse and multi-dimensional indicators. Costanza et al. (1997), Millennium Ecosystem Assessment (2005) and Fu et al. (2017) explored the supply-demand relationship of ecosystem services by constructing an indicator system. Other scholars have also conducted research and testing on the selection of ecosystem service supply and demand indicators. For example, Layke et al. (2012) graded different types of ecosystem services through standards such as information transmission capacity and data availability, and analyzed the strengths and weaknesses of each indicator by comparing the results. Van Oudenhoven et al. (2012) believed that the indicators used to assess the supply and demand of ecosystem services should be quantifiable, changeable on a spatiotemporal scale, and sensitive to land use changes.
In the development of ecosystem service supply and demand research, the importance of spatiotemporal scales has gradually become prominent. Since the production and ecosystem service spatial flows depend on ecological and geographic processes at different spatiotemporal scales, and involve different fields such as biology and society, there are scale effects in all aspects of the ecosystem service supply and demand simulation. For instance, there are significant spatial scale, ecological scale and policy scale effects on the supply capacity and benefit degree of the De Wieden wetland ecosystem services (Hein et al., 2006). In addition, some scholars have begun to explore the multi-scale characteristics of ecosystem service sensitivity under disturbance scenarios at different scales and to discuss the problem of the scale matching between ecosystem management measures and ecosystem services (Gabriel et al., 2010; Petrosillo et al., 2010).
More and more scholars are exploring the relationship between ecosystem services and human well-being by combining the ecosystem service supply and demand. However, most of these studies on spatial characteristics are still static. There are still relatively few clear research systems which aim at the ecosystem service dynamic trajectory during the spatial transfer progress and the distribution of benefit degrees in different demand areas.

3 Drivers of changes in the supply and demand of ecosystem services

Global change significantly affects the stability of ecosystems by changing their structures and functions, causing continuous changes in the supply and demand of ecosystem services, which seriously threatens the human living environment and the sustainable development of the social economy. As the two global issues most closely related to human activities, land use change and climate change are the two main factors affecting the temporal and spatial distributions and changes in ecosystem services.

3.1 The impact of land use changes on ecosystem services

As an important link between human activities and environmental feedback (Yu and Yang, 2002), land use and land cover changes are the primary aspects of global environmental change. Land is the basic structure of the ecosystem. Land use and its quantity, pattern and changes directly affect the pattern and process of the ecosystem in a specific area, and further affect the availability of ecosystem products and services. Thus, land use change has an impact on human society (Fu and Zhang, 2014; Shiferaw et al., 2019). Most studies suggest that the land use and land cover changes (LUCC) caused by human activities have been one of the main reasons for the weakening of the current global ecosystem service supply capacity and the decline of service value (Li, 1996; MEA, 2005; Rimal et al., 2019).
The types and areas of land cover directly affect the types and intensity of ecosystem services (Tan et al., 2020). For instance, arable lands support a strong food supply service, and weak regulating and cultural services, while forests provide strong regulation services, such as soil and water conservation services, and climate regulation services, but relatively weak provision services. Xie et al. (2006) and Huang et al. (2012) reported that the total value of ecosystem services is primarily caused by the decreasing of the ecological area due to urban expansion. Estoque and Murayama (2012) indicated that the decline in forest area was the main reason for the decline of ecosystem service value in Baguio City, the Philippines.
The impact of land use activities on the intensity of ecosystem services has a significant temporal and spatial dependence, and the disturbances to their functions are unbalanced. Chen et al. (2019) thought that the intensity of ecosystem services and land use in counties in China are significantly negatively correlated with each other, but this correlation is quite different between the temporal and spatial scales. Arowolo et al. (2018) and García-Llamas et al. (2019) believed that the impact of land use change on ecosystem services is different or even opposite at different scales. For example, the economic value of the overall ecosystem would increase due to arable land expansion in a short period of time, whereas this expansion would result in greater economic losses in the long-term due to the degradation of environmental quality and the decline of ecosystem services (Arowolo et al., 2018). Huq et al. (2019) showed that the increase of wetlands in southern Bangladesh after the rainy season and the expansion of agricultural land before the rainy season led to an increasing value of ecosystem services in the watershed, even though the total ecosystem services and economic value in the long run were still on a downward trend.
LUCC also affect the relationships between ecosystem services, leading to different degrees of change in the pattern and supply capacity of different kinds of ecosystem services. Lv and Cheng (2007) found that the reclamation of grassland for cultivated land enhanced the value of ecosystem products and services, while the value of regulation and support services for maintaining water and soil decreased. Watson et al. (2020) revealed that there is a trade-off between the ecosystem services provided by natural ecosystems and the ecosystem services related to land use. For a certain area, the overall value of ecosystem services is mainly determined by the synergy of various services. Zheng et al. (2020) conducted a simulation study on the value of ecosystem services in the Three-River Headwaters Region from 1990 to 2015, and found that synergistic effects dominated ecosystem services more than trade-off effects at multiple scales. Therefore, Sharma et al. (2019) emphasized the necessity of maintaining an appropriate proportion of land use to maintain the best supply of ecosystem services.
Many previous studies have proven that land cover has varying degrees of impact on ecosystem services. However, due to the lack of reliable ecosystem service value evaluation factors, the effects of the indirect, direct, and overall driving forces on ecosystem service changes are still largely unclear.

3.2 The impact of climate change on ecosystem services

Climate change is another key issue of global change, since it can directly or indirectly affect ecosystem services by changing the ecological structure and natural processes. Most studies have shown that climate change affects the supply capacity of most ecosystem services, and the vulnerability of ecosystem services is more significant under extreme weather conditions. The Millennium Ecosystem Assessment revealed that global climate change has led a decline in the supply capacity for most ecosystem services (MEA, 2005). In some areas, climate change was even considered to be a key factor in the decline of local ecosystem services (Alva-Basurto and Arias-González, 2014). For example, changes in the natural environment affected the species that depend on it for survival to a certain extent, which in turn affected the supply capabilities and values of ecosystem services related to these species. The rising temperature promoted the growth of jellyfish, causing an increase in their number, which weakened the value of local eco-tourism services (Pecl et al., 2017). The species in the American oceans that are fished are not adapting to the rise in temperature and have moved in the polar direction, which has reduced the food supply capacity of the American oceans (Pinsky and Fogarty, 2012). On the other hand, due to the strong interrelationship between ecosystems and climate change, species changes have also affected the availability of ecosystem services and had a certain degree of impact on the local climate. For example, Pires et al. (2018) believed that an increasing species diversity maintains the relative stability of ecosystem services, and reduces the frequency and impact of extreme weather. However, the impacts of climate change on ecosystem services are not all negative. Researchers have found that even though there is a strong correlation between temperature and ecosystem services, the relevant direction depends on the types of ecosystem services (Ding and Nunes, 2014).
Climate change also affects ecosystem service management. The spatial differences of climatic factors determine that the impacts of climate change on ecosystem services are regional. Therefore, analyzing the responses of ecosystem services to climate change in a region/local area can provide a scientific basis for the management of the ecosystem services in that region. At present, some scholars have simulated the impact of climate change on different ecosystem services and vulnerabilities in Australia, Europe and the United States through scenario simulation or by establishing a conceptual framework, which can be used to explore the strategies and methods for the sustainable management of natural resources, laying a foundation for the sustainable development of those resources (Metzger and Schröter, 2006; Bryan, 2011; Weiskopf et al., 2020). However, due to limitations in the data availability and the research scales, there have been relatively few similar studies on assessing the impact of climate change on various ecosystem services at the national scale. There is still a lack of systematic research on the comprehensive impact of climate change on various ecosystem services.
In other words, climate change has mostly negative impacts on ecosystem services, but the specific impacts vary with the different driving factors, types of services evaluated, and research methods (Runting et al., 2017). Temperature, precipitation, and sea level rise are the direct drivers most commonly selected by researchers for exploring changes in ecosystem service supply. In addition, due to the high complexity and spatial differences of climate change, the demand side of ecosystem services is usually not considered. Therefore, the results also have a certain degree of uncertainty, which creates a certain degree of difficulty for the management of ecosystems and natural resources.
Changes in ecosystem processes are often very complicated. Multiple driving factors affect the state and conditions of the ecosystem at the same time. In addition, the function of the ecosystem changes through self-adaptation and other methods, which ultimately leads to dynamic changes in ecosystem services. Therefore, research on the changes in ecosystem services needs to integrate various driving factors such as ecosystem structure, biodiversity, human cognition, economic development level, and social ecological knowledge level (Li et al., 2011; Zhao et al., 2018). Direct driving factors such as land cover change and climate change have a significant adverse impact on ecosystem services; while indirect driving factors including population, economy, politics, science and technology have more scattered impacts on the ecosystem, which are generally recognized by the current viewpoint (Norgaard, 2010; Robards et al., 2011). However, most of the causal relationships between ecosystem services and the factors driving their changes are based on scenarios of future land cover or climate changes in the ecosystem (Nelson et al., 2010; Lawler et al., 2014; Rosenzweig et al., 2014; Martinez-Harms et al., 2017). Few studies have evaluated the impacts of drivers on changing trends in ecosystem services (Nelson et al., 2013; Schirpke et al., 2013; Guerra et al., 2016; Egarter et al., 2016). This limitation hinders the relevant research on the ecosystem service driving mechanisms and affects the precise implementation of landscape planning and protection policies.

4 Research progress on the ecosystem service spatial flow

The research related to ecosystem service spatial flow is currently still in the preliminary stage. At this early stage, the concept of ecosystem service flow has been the main discussion content. Some scholars emphasize the spatial flow process, believing that ecosystem service flow should be a spatial dynamic transport progress from the supply area to the beneficiary area (Syrbe and Walz, 2012; Xiao et al., 2017), while other scholars emphasize the ultimate utility of ecosystem services. Mostly based on human needs, this explanation often suggests that the flow of ecosystem services only includes the energy flow consumed by the ecosystem itself and the part used by humans, without considering the state and structure of the ecosystem (Pulselli et al., 2011). On the whole, ecosystem service flow is based on the relationship between service supply and demand, driven by nature and humans, transferring from the supply sources to the benefit sinks in the spatiotemporal scale. Although the classification system and conceptual interpretation of the definition of the ecosystem service flow are relatively mature, the specific methods for quantifying and spatially mapping it are rarely mentioned.
With the extensive discussion of the concept, the temporal and spatial characteristics, carrier characteristics, and quantitative attributes of ecosystem service spatial flows are generally recognized. Li et al. (2014) and Johnson et al. (2012) indicated that ecosystem services must rely on a certain form which could carry them from the supply area to the beneficiary area. Therefore, the quantity, direction, and speed based on the carriers are the three main aspects when simulating and quantifying the spatial flow of ecosystem services. Moreover, since some carriers are a form of ecosystem services themselves, and even affect multiple services at the same time, there is a complex correspondence between carriers and ecosystem services. The same carrier could correspond to a variety of ecosystem service flows, and an ecosystem service flow could also be affected by multiple carriers.
In the current research on the spatial flow of ecosystem services, many scholars recognize the importance of simulating and quantifying the process of ecosystem services from supply unit to benefit unit, and simulate the flow process of ecosystem services by establishing a conceptual framework. For example, Hou et al. (2020) constructed a conceptual framework based on the socio-economic cascade diagram of the ecosystem to explore the relationship between the nutrient regulation service flow, demand and potential supply capacity. Shi et al. (2020) used linear programming and simplex methods to simulate the flow of food supply, carbon fixation and oxygen release, leisure and entertainment, and biological habitat protection services in Shanghai. Vallecillo et al. (2019) used the experimental ecosystem accounting (SEEA EEA) evaluation framework to identify and evaluate the flow of entertainment services, service potential capacity, and the status of supply and demand in European natural landscapes. From the perspective of significant spatial heterogeneity of ecosystem services, Schirpke et al. (2019) used an evaluation framework to evaluate the flow directions and specific transfer mechanisms of six main ecosystem services in the Alps and surrounding lowlands at the regional and global levels. Matthias Schröter et al. (2018) showed the moving trajectory and mechanism of the coffee trade, migration of northern tail ducks, flood regulation in the Danube Basin, and information flow of giant pandas by constructing a conceptual framework of interregional ecosystem service flow. Through this method, the ecosystem services obtained by the beneficiary areas that come from the flow of services could be quantified clearly, but it is unable to give the exact path of ecosystem services or to simulate their spatial flow process.
The spatially distributed model emphasizes the service flow process, so it can simulate the specific transfer trajectory, direction and quantity of ecosystem services from the supply area to the demand area. Investigators at the University of Vermont in the United States carried out research on the Artificial Intelligence for Ecosystem Services (ARIES) project with funding from the National Natural Science Foundation of the United States. At present, the ARIES model has developed eight modules for the simulation of ecosystem service spatial flow, including flood regulation, fresh water supply, carbon sinks and carbon storage, which has been widely used in various regions at home and abroad. For example, Li et al. (2017) used the InVEST and ARIES systems to simulate the flow of freshwater regulation services from supply to consumption on different spatiotemporal scales in the Beijing-Tianjin-Hebei region. Bagstad et al. (2014) quantified the supply and demand paths and flow processes of five services in Puget Sound, Washington, USA, through the methodology of ARIES. On the basis of that study, Zank et al. (2016) studied the impact of urban expansion on natural resource storage and ecosystem service flows.
In addition, Bagstad et al. (2013) proposed a “Service Path Attribution Networks” (SPANs) model by comparing the ARIES and InVEST models. This model, which integrated various simulation models that are commonly used in ecology and geography, uses probabilistic Bayesian networks to analyze the flow of ecosystem services from supply locations to benefit areas, such as simulating the flow paths of trade commodities, passive biophysical flows, species migration and diffusion, and information flow (Koellner et al., 2019). The SPANs model simulates the specific flow path of ecosystem services, which makes up for the lack of a systematic approach and lays a foundation for future research on simulating the spatial flow and revealing the theory and actual spatial transfer process of ecosystem services. However, because the ecosystem service delivery process is much more complicated, and the spatially distributed model requires a large amount of data for the calculations, the simulation results also have certain deviations.
So far, most of the conceptual research on ecosystem service flow has focused on the potential supply, while the considerations of service demand and the flow trajectory are quite few. Cultural services, involving non-material services such as spiritual level, cognitive level, and aesthetic experience which are usually hard to quantify (Shaw et al., 2016; Tenerelli et al., 2016), are now of less concern to scholars. Although the diversity of ecosystem service flows on the spatial scale are more often recognized by scholars (Boyd and Banzhaf, 2007), the temporal scale characteristics are generally ignored (Fisher et al., 2009; Ruhl et al., 2013). Therefore, in future research, more attention should be paid to the complete process of the ecosystem service spatial flow at a range of spatiotemporal scales under the trend of quantitative analysis and model simulations.

5 Conclusions

Assessing the balance of the supply and demand of ecosystem services can provide a scientific foundation for landscape planning and ecosystem management, which can support an important guarantor for the regional ecological security as well. Exploring the driving factors of ecosystem services is conducive to predicting the trends of ecosystem service changes so that management measures can be adjusted in time. Simulating the real status of the spatial flow of ecosystem services can clarify the transmission process from generation to use, establish a feedback relationship between ecosystem service supply and demand, and also provide references for formulating ecological compensation and other policy management measures.
Although some achievements have been made in assessing the relationships between the supply and demand of ecosystem services, there are still some shortcomings. Firstly, there are very few cultural service indicators that can be quantified easily, and the carrier form of these services is relatively abstract. Thus, compared to provision services, regulation services and support services, there are relatively few studies on cultural services. Secondly, a number of studies have simulated the spatial flow paths of ecosystem services. However, most of them are still based on static assessments according to the relationship between supply and demand. The dynamic studies are relatively few at this point. As a result, quantifying the flow and direction of regional ecosystem services, and figuring out the rules of the spatiotemporal distribution in the process of ecosystem service transfer still need more in-depth study.
In addition, there are still some issues which need to be explored in the future. The entire process of ecosystem services from supply to demand is affected by both nature and human society. Quantifying the influences of various driving factors and revealing the specific driving mechanisms of different ecosystem services are critical for promoting theoretical research on ecosystem services and spatial flow processes in the future. The prediction of ecosystem services and their values is another issue that needs to be studied in the future as well. In addition, for the formulation of ecosystem management policies, it is important to analyze the changing trends of ecosystem services under different scenarios on the basis of a clear driving mechanism.
Finally, in the research on ecosystem services, the spatio-temporal scale is a very important research basis. Most studies have fully taken into account the boundaries of the spatial and temporal scales. However, the analysis of driving factors usually considers the influence of the time scale, while the analysis of spatial flow focuses more on the spatial scale. In addition, the spatial distribution, flow status, and benefits of the same ecosystem services all depend on the various spatiotemporal scales due to the spatial heterogeneity. Hence, it is also necessary to study the conversion of different spatiotemporal scales of ecosystem services and its spatial flows.

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