Ze-Xin Fan

Leaf development and photosynthetic properties of three tropical tree species with delayed greening

Leaf developmental patterns were characterized for three tropical tree species with delayed greening. Changes in the pigment contents, photosynthetic capacity, stomata development, photosystem 2 efficiency, rate of energy dissipation, and the activity of partial protective enzymes were followed in developing leaves in an attempt to elucidate the relative importance of various photoprotective mechanisms during leaf ontogeny. Big leaves of Anthocephalus chinensis, a fast-growing light demanding species, expanded following an exponential pattern, while relatively small leaves of two shade-tolerant species Litsea pierrei and Litsea dilleniifolia followed a sigmoidal pattern. The juvenile leaves of A. chinensis and L. pierrei contained anthocyanin located below the upper epidermis, while L. dilleniifolia did not contain anthocyanin. Leaves of A. chinensis required about 12 d for full leaf expansion (FLE) and photosynthetic development was delayed 4 d, while L. pierrei and L. dilleniifolia required 18 or 25 d for FLE and photosynthetic development was delayed 10 or 15 d, respectively. During the leaf development the increase in maximum net photosynthetic rate was significantly related to changes in stomatal conductance and the leaf maturation period was positively related to the steady-state leaf dry mass per area for the three studied species. Dark respiration rate of leaves at developing stages was greater, and pre-dawn initial photochemical efficiency was lower than that of mature leaves. Young leaves displayed greater energy dissipation than mature leaves, but nevertheless, the diurnal photoinhibition of young L. dilleniifolia leaves was higher than that of mature leaves. The young red leaves of A. chinensis and L. pierrei with high anthocyanin contents and similar diurnal photoinhibition contained more protective enzymes (superoxide dismutase, ascorbate peroxidase) than mature leaves. Consequently, red leaves may have higher antioxidant ability.

Water-use advantage for lianas over trees in tropical seasonal forests

Lianas exhibit peak abundance in tropical forests with strong seasonal droughts, the eco-physiological mechanisms associated with lianas coping with water deficits are poorly understood. We examined soil water partitioning, sap flow, and canopy eco-physiological properties for 99 individuals of 15 liana and 34 co-occurring tree species in three tropical forests that differed in soil water availability. In the dry season, lianas used a higher proportion of deep soil water in the karst forest (KF; an area with severe seasonal soil water deficit (SSWD)) and in the tropical seasonal forest (TSF, moderate SSWD), permitting them to maintain a comparable leaf water status than trees in the TSF or a better status than trees in the KF. Lianas exhibited strong stomatal control to maximize carbon fixation while minimizing dry season water loss. During the dry period, lianas significantly decreased water consumption in the TSF and the KF. Additionally, lianas had a much higher maximum photosynthetic rates and sap flux density in the wet season and a lower proportional decline in photosynthesis in the dry season compared with those of trees. Our results indicated that access to deep soil water and strong physiological adjustments in the dry season together with active wet-season photosynthesis may explain the high abundance of lianas in seasonally dry forests.

Spatial and temporal patterns of variations in tree growth over the northeastern Tibetan Plateau during the period AD 1450–2001

We analyzed spatial and temporal growth variations of Qilian Junipers over the northeastern Tibetan Plateau (TP) during the period 1450—2001 by applying the empirical orthogonal function (EOF) technique to seven moisture-sensitive tree ring-width chronologies. The first three principal components together explain 78.21% of the total variance. The first principal component (PC1) accounts for 51.58% of the total variance, and represents consistent tree-growth variations in the northeastern TP. The second EOF mode reveals an opposite structure for the southern and northern parts of the northeastern Tibetan Plateau, reflecting the effect of huge topography and associated atmospheric circulation on tree-growth anomalies. The third EOF mode represents an opposite structure for the eastern and western parts of the northeastern Tibetan Plateau, possibly indicating the influence of different branches of atmospheric circulation system in this area. The EOF analysis results of instrumental precipitation data resembl...We analyzed spatial and temporal growth variations of Qilian Junipers over the northeastern Tibetan Plateau (TP) during the period 1450—2001 by applying the empirical orthogonal function (EOF) technique to seven moisture-sensitive tree ring-width chronologies. The first three principal components together explain 78.21% of the total variance. The first principal component (PC1) accounts for 51.58% of the total variance, and represents consistent tree-growth variations in the northeastern TP. The second EOF mode reveals an opposite structure for the southern and northern parts of the northeastern Tibetan Plateau, reflecting the effect of huge topography and associated atmospheric circulation on tree-growth anomalies. The third EOF mode represents an opposite structure for the eastern and western parts of the northeastern Tibetan Plateau, possibly indicating the influence of different branches of atmospheric circulation system in this area. The EOF analysis results of instrumental precipitation data resemble those derived from the seven regional chronologies, providing further support that tree growth is a good indicator of regional precipitation variations. According to the PC1, dry conditions occurred in AD 1450—1510, 1631—1739, 1765—1833, 1866—1883 and 1921—1947, whereas periods of relatively wet years are identified for AD 1511—1630, 1740—1764, 1834—1865, 1884—1920 and 1948—2001. During the past 550 years, 28 years with extreme regional growth depressions was identified. These ring-width extremes generally occurred in century-scale dry periods. The years 1467, 1484, 1641, 1721, and 1928 with extreme tree-growth reductions in the northeastern TP are coeval with severe drought events in eastern China. In addition, the dry intervals or extreme growth depressions generally coincided with cold periods in the northeastern TP and vice versa, implying that the probability of occurrence of wet events in the northeastern TP will be higher in the future under the ongoing global warming.

Continental-Scale Temperature Variability during the Past Two Millennia: Supplementary Information

Past global climate changes had strong regional expression. To elucidate their spatio-temporal pattern, we reconstructed past temperatures for seven continental-scale regions during the past one to two millennia. The most coherent feature in nearly all of the regional temperature reconstructions is a long-term cooling trend, which ended late in the nineteenth century. At multi-decadal to centennial scales, temperature variability shows distinctly different regional patterns, with more similarity within each hemisphere than between them. There were no globally synchronous multi-decadal warm or cold intervals that define a worldwide Medieval Warm Period or Little Ice Age, but all reconstructions show generally cold conditions between ad 1580 and 1880, punctuated in some regions by warm decades during the eighteenth century. The transition to these colder conditions occurred earlier in the Arctic, Europe and Asia than in North America or the Southern Hemisphere regions. Recent warming reversed the long-term cooling; during the period ad 1971–2000, the area-weighted average reconstructed temperature was higher than any other time in nearly 1,400 years.

Radial Growth of Qilian Juniper on the Northeast Tibetan Plateau and Potential Climate Associations

There is controversy regarding the limiting climatic factor for tree radial growth at the alpine treeline on the northeastern Tibetan Plateau. In this study, we collected 594 increment cores from 331 trees, grouped within four altitude belts spanning the range 3550 to 4020 m.a.s.l. on a single hillside. We have developed four equivalent ring-width chronologies and shown that there are no significant differences in their growth-climate responses during 1956 to 2011 or in their longer-term growth patterns during the period AD 1110–2011. The main climate influence on radial growth is shown to be precipitation variability. Missing ring analysis shows that tree radial growth at the uppermost treeline location is more sensitive to climate variation than that at other elevations, and poor tree radial growth is particularly linked to the occurrence of serious drought events. Hence water limitation, rather than temperature stress, plays the pivotal role in controlling the radial growth of Sabina przewalskii Kom. at the treeline in this region. This finding contradicts any generalisation that tree-ring chronologies from high-elevation treeline environments are mostly indicators of temperature changes.

Multi-proxy temperature reconstruction from the West Qinling Mountains, China for the past 500 years.

A total of 290 tree-ring samples, collected from six sites in the West Qinling Mountains of China, were used to develop six new standard tree-ring chronologies. In addition, 73 proxy records were assembled in collaboration with Chinese and international scholars, from 27 publically available proxy records and 40 tree-ring chronologies that are not available in public datasets. These records were used to reconstruct annual mean temperature variability in the West Qinling Mountains over the past 500 years (AD 1500–1995), using a modified point-by-point regression (hybrid PPR) method. The results demonstrate that the hybrid PPR method successfully integrates the temperature signals from different types of proxies, and that the method preserves a high degree of low-frequency variability. The reconstruction shows greater temperature variability in the West Qinling Mountains than has been found in previous studies. Our temperature reconstruction for this region shows: 1) five distinct cold periods, at approximately AD 1520–1535, AD 1560–1575, AD 1610–1620, AD 1850–1875 and AD 1965–1985, and four warm periods, at approximately AD 1645–1660, AD 1705–1725, AD 1785–1795 and AD 1920–1945; 2) that in this region, the 20th century was not the warmest period of the past 500 years; and 3) that a dominant and persistent oscillation of ca. 64 years is significantly identified in the 1640–1790 period.

MAY-JUNE MEAN TEMPERATURE RECONSTRUCTION OVER THE PAST 300 YEARS BASED ON TREE RINGS IN THE QILIAN MOUNTAINS OF THE NORTHEASTERN TIBETAN PLATEAU

A juniper (Juniperus przewalskii Kom.; synonym: Sabina przewalskii) tree-ring width chronology was developed to investigate the regional climate variability for the Qilian Mountains. Statistically, the chronology was appropriate for reconstructing the regional mean temperature of May-June from A.D.1700 to the present. The phenomenon of synchronous extremely high temperatures and extreme droughts in the 1920s was revealed by comparing our reconstruction with drought events in this region. Multi-taper spectral analysis indicated the existence of significant low- and high-frequency periods (40–46 years, 34, 23–25, 5.6, 2.1, 2.5–2.8 years). Overall, the study not only extended the temperature record, but also provided reliable long-term temperature information to help understand the possible forcing of climate changes in the Qilian Mountains.

Time lags between crown and basal sap flows in tropical lianas and co-occurring trees

Water storage in the stems of woody plants contributes to their responses to short-term water shortages. To estimate the contribution of water storage to the daily water budget of trees, time lags of sap flow between different positions of trunk are used as a proxy of stem water storage. In lianas, another large group of woody species, it has rarely been studied whether stored water functions in their daily water use, despite their increasing roles in the carbon and water dynamics of tropical forests caused by their increasing abundance. We hypothesized that lianas would exhibit large time lags due to their extremely long stems, wide vessels and large volume of parenchyma in the stem. We examined time lags in sap flow, diel changes of stem volumetric water content (VWC) and biophysical properties of sapwood of 19 lianas and 26 co-occurring trees from 27 species in 4 forests (karst, tropical seasonal, flood plain and savanna) during a wet season. The plants varied in height/length from <5 to >60 m. The results showed that lianas had significantly higher saturated water content (SWC) and much lower wood density than trees. Seven of 19 liana individuals had no time lags; in contrast, only 3 of 26 tree individuals had no time lags. In general, lianas had shorter time lags than trees in our data set, but this difference was not significant for our most conservative analyses. Across trees and lianas, time lag duration increased with diurnal maximum changeable VWC but was independent of the body size, path length, wood density and SWC. The results suggest that in most lianas, internal stem water storage contributes little to daily water budget, while trees may rely more on stored water in the stem.

Analysis of reconstructed annual precipitation from tree-rings for the past 500 years in the middle Qilian Mountain

The ring-width chronology of a Juniperus przewalskii tree from the middle of the Qilian Mountain was constructed to estimate the annual precipitation (from previous August to current July) since AD 1480. The reconstruction showed four major alternations of drying and wetting over the past 521 years. The rainy 16th century was followed by persistent drought in the 17th century. Moreover, relatively wet conditions persisted from the 18th to the beginning of 20th century until the recurrence of a drought during the 1920s and 1930s. Based on the Empirical Mode Decomposition method, eight Intrinsic Mode Functions (IMFs) were extracted, each representing unique fluctuations of the reconstructed precipitation in the time-frequency domain. The high amplitudes of IMFs on different timescales were often consistent with the high amount of precipitation, and vice versa. The IMF of the lowest frequency indicated that the precipitation has undergone a slow increasing trend over the past 521 years. The 2–3 year and 5–8 year time-scales reflected the characteristics of inter-annual variability in precipitation relevant to regional atmospheric circulation and the El Niño-Southern Oscillation (ENSO), respectively. The 10–13 year scale of IMF may be associated with changing solar activity. Specifically, an amalgamation of previous and present data showed that droughts were likely to be a historically persistent feature of the Earth’s climate, whereas the probability of intensified rainfall events seemed to increase during the course of the 19th and 20th centuries. These changing characteristics in precipitation indicate an unprecedented alteration of the hydrological cycle, with unknown future amplitude. Our reconstruction complements existing information on past precipitation changes in the Qilian Mountain, and provides additional low-frequency information

The Cooling Trend of Canopy Temperature During the Maturation, Succession, and Recovery of Ecosystems

The maximum exergy dissipation theory provides a theoretical basis for using surface temperature to measure the status and development of ecosystems, which could provide an early warning of rapid evaluation of ecosystem degradation. In the present study, we used the radiation balance of ecosystems to demonstrate this hypothesis theoretically. Further, we used empirical data to verify whether ecosystems gain more radiation, while lowering their surface temperatures, as they develop naturally. We analyzed 12 chronosequences from the FLUXNET database using meteorological data and heat fluxes. We included age, disturbance, and successional chronosequences across six climate zones. Net radiation (Rn) and the ratio of net radiation to global radiation (Rn/Rg) were used to measure the energy gain of the ecosystems. The maximum daily air temperature above the canopy (Tmax) and thermal response number (TRN) were used to analyze the surface temperature trends with ecosystem natural development. The general trends of Tmax, TRN, Rn, and Rn/Rg demonstrated that ecosystems become cooler and more stable, yet gain more energy, throughout their natural development. Among the four indicators, TRN showed the most consistent trends and highest sensitivity to ecosystem growth, succession, and recovery. Moreover, TRN was not significantly influenced by precipitation or wind. We propose that TRN can be used to rapidly evaluate or warn of ecosystem disturbance, senescence, and degradation without prior knowledge of species composition, nutrient status, and complex ecosystem processes.