1. Introduction
Plant phenology is a mutually adaptive growth and development rhythm formed by long-term adaptation of plants to seasonal changes in environmental conditions such as temperature, precipitation, and light [
1], which not only reflects plant growth and development but also indicates climate change [
2] and has been widely used to guide agricultural activities and disaster prevention and mitigation [
3]. Meanwhile, plant phenology affects water-heat exchange and carbon cycling in ecosystems [
4,
5], and is an important parameter in land surface process models and plant productivity models [
6]. Therefore, it is important to study plant phenology patterns and their relationship with environmental conditions.
It was found that plant spring phenology tends to be earlier and autumn phenology tends to be later in a warming context [
7,
8], and this phenomenon was more pronounced in the high latitudes of the Northern Hemisphere [
9]. However, there is a clear spatial variability in the response of grassland plant phenology to climate change in Inner Mongolia, with a trend of earlier green-up and later wilting in the southern region, and a trend of later green-up and earlier wilting in the central and northern regions [
10], and the factors causing spatial variability in grassland plant phenology are still unclear. Meteorological factors that affect plant phenology include temperature, precipitation, light, air humidity, carbon dioxide concentration, etc. Among them, temperature is considered to be the most important environmental factor affecting plant phenology [
11], and plants can only grow and develop in a certain temperature environment and need a certain cumulative temperature to complete their life cycle [
12]. Water deficit limits the use of light and heat conditions by plants [
13] and is considered a key factor in regulating vegetation activity in arid and semi-arid regions [
14]. Light is the source of energy for photosynthesis in plants, and the organic matter produced by photosynthesis is the material basis for plant growth and development [
15], and photoperiod has also been shown to be an important indicator of how light affects plant phenology [
16]. Meanwhile, there are significant interactions between temperature, precipitation, and light, which together affect the changes in plant phenology [
17,
18]. Phenology models have evolved from statistical to mechanistic models, and have been widely used for simulation of phenology and prediction of future climate change impacts in areas lacking phenology observations [
19]. However, limited by the understanding of the mechanisms influencing phenological changes, especially because most of the existing studies only consider the phenological effects from single or several environmental factors on specific plant species, the response of plant phenology to the combined effects of environmental factors and its mechanisms are still unclear, making these models are not able to reflect the realism of vegetation growth and cannot effectively simulate phenological periods [
20].
The research showed that plant phenology was closely related to the dynamics of photosynthesis [
21]. Plant photosynthesis is the result of the interaction between environmental factors and plant biological properties, reflecting the influence of the total climate production factors. Climate is the most important factor affecting plant growth and development and is the basis for morphological establishment and physiological and biochemical changes in plants [
22,
23]. Climatic production potential refers to the highest biological or agricultural yield per unit area of land when other conditions (e.g., soil, nutrients, carbon dioxide, etc.) are at optimum conditions and the local climatic resources such as light, heat, and water are fully and rationally utilized [
24]. Climate production potential not only reflects the influence of climate factors (temperature, moisture, light, etc.) and their combined effects on plant production, but also ensures the uniformity of the influencing factors during the whole process of plant growth and the cyclical changes in their interaction with the environment; at the same time, climate production potential also reflects the combined effects of biological factors (e.g., leaf area), environmental factors and their interactions, and can reflect the effects of extreme weather and climate events. Therefore, the use of climate production potential as a driver of plant phenology changes can avoid the shortcomings of existing models and achieve accurate simulation of phenology [
25].
Chinese temperate grasslands are the third largest in the world [
26], sensitive to climate change, and play an important role in the global carbon cycle [
27]. The grassland of
S. krylovii is one of the representative types of typical grasslands [
28], which occupies an important position in livestock production [
29] and has been significantly affected by warm and dry climate [
30]. Using the long-term phenological and corresponding meteorological observation data from 1985 to 2018 in the grassland of
S. krylovii, this study proposes the new concepts of climate production potential, cumulative climate production potential (reflecting resource accumulation), first-order derivative of cumulative climate production potential (reflecting the rate of resource change) and second-order derivative of cumulative climate production potential (reflecting sudden resource change) based on the total climate production factors influencing plant phenological changes [
25]. On this basis, this study will intend to (1) verify the feasibility of simulating plant phenology and triggering thresholds based on total climatic production factors, and (2) clarify the relationship between the main phenological periods of
S. krylovii and total climatic production factors and their triggering thresholds to improve the understanding of the response of phenology to the combined effect of meteorological conditions and provide a basis for the development of phenological models.
4. Discussion and Conclusion
Meteorological factors are important factors that affect plant growth and development, as well as the basis for plant morphogenesis, physiological and biochemical changes. The meteorological factors affecting the green-up period of grassland plants include temperature, sunshine duration [
36], accumulated temperature [
19,
37] and moisture [
38,
39]. When the air temperature is below the threshold, the autumn phenology occurs [
40]; when the photoperiod is shortened to the threshold that limits plant growth and development, it will induce plant leaf senescence and enter a dormant state. The in situ simulation experiment of
S. krylovii grassland also showed that it is environmental factors rather than plant productivity that drives leaf senescence [
42]. In this study, the abrupt relationship between the green-up date and wilting date of
S. krylovii plant and the second derivative of cumulative climate production potential was the best, indicating that different phenological periods of grassland plants were comprehensively affected by meteorological conditions, and sudden changes in environmental factors trigger the green-up and wilting of plants. The heading date of
S. krylovii plants had the best relationship with the cumulative climate production potential, indirectly proving the impact of rising summer temperature as a driver of climate production potential on the phenological periods between the beginning and ending phenology, reflecting the cumulative climate resource effects.
In this study, the long-term phenology and corresponding meteorological observation data of S. krylovii grassland from 1985 to 2018 were used to verify the indication of climate production potential based on total climate production factors on plant phenology, and the relationship between the main phenological stages of S. krylovii plant and climate production potential was clarified. The climate production potential not only reflects the comprehensive effect of environmental factors in plant production and their consistency throughout the entire process of plant growth, but also reflects the effects of biological factors, environmental factors and their interactions, as well as extreme weather and climate events, which will be helpful for achieving accurate simulation of plant phenology. The main conclusions are as following:
(1) From 1985 to 2018, the green-up date of
S. krylovii plant in Xilinhot, Inner Mongolia was delayed by an average of 5.4 d/10a, the heading date was advanced by an average of 5.4 d/10a, the wilting date was advanced by an average of 1.2 d/10a, and the length of the growing season was shortened by an average of 6.3 d/10a. The phenological change trend of
S. krylovii is consistent with the existing research in this area [
36,
43].
(2) The climate production potential based on total climate production factors is a good indicator for plant phenology, and the sudden change in cumulative climate production potential reflects the drastic changes in climate conditions, which can effectively indicate the green-up and wilting periods of S. krylovii plant; while the cumulative climate production potential reflects the rate of climate resource change, and can reflects resource utilization to a certain degree, which can effectively indicate the heading period of S. krylovii plant.
(3) The relationship between the phenological periods and the cumulative climate production potential of S. krylovii plant further indicates that different phenological periods of grassland plants are comprehensively influenced by meteorological conditions. To comprehensively understand the response of plant phenology to environmental changes, it is necessary to understand the influencing mechanism of total climate production factor on plant phenology.