Regulatory Mechanisms of Heat Stress Response and Thermomorphogenesis in Plants.

As worldwide warming intensifies, the average temperature of the earth continues to increase. Temperature is a key factor for the growth and development of all organisms and governs the distribution and seasonal behavior of plants. High temperatures lead to various biochemical, physiological, and morphological changes in plants and threaten plant productivity. As sessile organisms, plants are subjected to various hostile environmental factors and forced to change their cellular state and morphological architecture to successfully deal with the damage they suffer. Therefore, plants have evolved multiple strategies to cope with an abnormal rise in temperature. There are two main mechanisms by which plants respond to elevated environmental temperatures. One is the heat stress response, which is activated under extremely high temperatures; the other is the thermomorphogenesis response, which is activated under moderately elevated temperatures, below the heat-stress range. In this review, we summarize recent progress in the study of these two important heat-responsive molecular regulatory pathways mediated, respectively, by the Heat Shock Transcription Factor (HSF)-Heat Shock Protein (HSP) pathway and PHYTOCHROME INTER-ACTING FACTOR 4 (PIF4) pathways in plants and elucidate the regulatory mechanisms of the genes involved in these pathways to provide comprehensive data for researchers studying the heat response. We also discuss future perspectives in this field.

Ethylene Response Factor LlERF110 Mediates Heat Stress Response via Regulation of LlHsfA3A Expression and Interaction with LlHsfA2 in Lilies (Lilium longiflorum).

Heat stress seriously affects the quality of cut lily flowers. The ethylene response factors (ERFs) participate in heat stress response in many plants. In this study, heat treatment increased the production of ethylene in lily leaves, and exogenous ethylene treatment enhanced the heat resistance of lilies. LlERF110, an important transcription factor in the ethylene signaling pathway, was found in the high-temperature transcriptome. The coding region of LlERF110 (969 bp) encodes 322 amino acids and LlERF110 contains an AP2/ERF typical domain belonging to the ERF subfamily group X. LlERF110 was induced by ethylene and was expressed constitutively in all tissues. LlERF110 is localized in the nucleus and has transactivation activity. Virus-induced gene silencing of LlERF110 in lilies reduced the basal thermotolerance phenotypes and significantly decreased the expression of genes involved in the HSF-HSP pathway, such as LlHsfA2, LlHsfA3A, and LlHsfA5, which may activate other heat stress response genes; and LlHsp17.6 and LlHsp22, which may protect proteins under heat stress. LlERF110 could directly bind to the promoter of LlHsfA3A and activate its expression according to the yeast one hybrid and dual-luciferase reporter assays. LlERF110 interacts with LlHsfA2 in the nucleus according to BiFC and the yeast two-hybrid assays. In conclusion, these results indicate that LlERF110 plays an important role in the basal thermotolerance of lilies via regulation of the HSF-HSP pathway, which could be the junction of the heat stress response pathway and the ethylene signaling pathway.

Small HSPs play an important role in crosstalk between HSF-HSP and ROS pathways in heat stress response through transcriptomic analysis in lilies (Lilium longiflorum).

BACKGROUND: High temperature seriously limits the annual production of fresh cut lilies, which is one of the four major cut flowers in the global cut flower market. There were few transcriptomes focused on the gene expression of lilies under heat stress. In order to reveal the potential heat response patterns in bulbous plants and provide important genes for further genetic engineering techniques to improve thermotolerance of lily, RNA sequencing of lilies under heat treatments were conducted. RESULTS: In this study, seedlings of Lilium longiflorum 'White Heaven' were heat-treated at 37 °C for different lengths of time (0 h, 0.5 h, 1 h, 3 h, 6 h, and 12 h with a 12 h-light/12 h-dark cycle). The leaves of these lily seedlings were immediately collected after heat treatments and quickly put into liquid nitrogen for RNA sequencing. 109,364,486-171,487,430 clean reads and 55,044 unigenes including 21,608 differentially expressed genes (DEGs) (fold change ≥2) were obtained after heat treatment. The number of DEGs increased sharply during the heat treatments of 0.5 h-1 h and 1 h-3 h compared to that of other periods. Genes of the heat stress transcription factor (HSF) family and the small heat shock proteins (small HSPs, also known as HSP20) family responded to heat stress early and quickly. Compared to that of the calcium signal and hormone pathways, DEGs of the HSF-HSP pathway and reactive oxygen species (ROS) pathway were significantly and highly induced. Moreover, they had the similar expression pattern in response to heat stress. Small HSPs family genes were the major components in the 50 most highly induced genes at each heat stress treatment and involved in ROS pathway in the rapid response to heat stress. Furthermore, the barley stripe mosaic virus induced gene silencing (BSMV-VIGS) of LlHsfA2 caused a significantly reduced thermotolerance phenotype in Lilium longiflorum 'White Heaven', meanwhile decreasing the expression of small HSPs family genes and increasing the ROS scavenging enzyme ascorbate peroxidase (APX) genes, indicating the potential interplay between these two pathways. CONCLUSIONS: Based on our transcriptomic analysis, we provide a new finding that small HSPs play important roles in crosstalk between HSF-HSP and ROS pathways in heat stress response of lily, which also supply the groundwork for understanding the mechanism of heat stress in bulbous plants.

The Heat Stress Transcription Factor LlHsfA4 Enhanced Basic Thermotolerance through Regulating ROS Metabolism in Lilies (Lilium Longiflorum).

Heat stress severely affects the annual agricultural production. Heat stress transcription factors (HSFs) represent a critical regulatory juncture in the heat stress response (HSR) of plants. The HsfA1-dependent pathway has been explored well, but the regulatory mechanism of the HsfA1-independent pathway is still under-investigated. In the present research, HsfA4, an important gene of the HsfA1-independent pathway, was isolated from lilies (Lilium longiflorum) using the RACE method, which encodes 435 amino acids. LlHsfA4 contains a typical domain of HSFs and belongs to the HSF A4 family, according to homology comparisons and phylogenetic analysis. LlHsfA4 was mainly expressed in leaves and was induced by heat stress and H2O2 using qRT-PCR and GUS staining in transgenic Arabidopsis. LlHsfA4 had transactivation activity and was located in the nucleus and cytoplasm through a yeast one hybrid system and through transient expression in lily protoplasts. Over expressing LlHsfA4 in Arabidopsis enhanced its basic thermotolerance, but acquired thermotolerance was not achieved. Further research found that heat stress could increase H2O2 content in lily leaves and reduced H2O2 accumulation in transgenic plants, which was consistent with the up-regulation of HSR downstream genes such as Heat stress proteins (HSPs), Galactinol synthase1 (GolS1), WRKY DNA binding protein 30 (WRKY30), Zinc finger of Arabidopsis thaliana 6 (ZAT6) and the ROS-scavenging enzyme Ascorbate peroxidase 2 (APX2). In conclusion, these results indicate that LlHsfA4 plays important roles in heat stress response through regulating the ROS metabolism in lilies.

Research on the dissipation of green tide and its influencing factors in the Yellow Sea based on Google Earth Engine.

Since 2007, the outbreak of green tides has become the most serious ecological problem in the Yellow Sea. In this study, a new method was used to identify green tides in multi-source satellite data from 2007 to 2020, and the relationship between the conditions necessary for green tide dissipation and other environmental factors was discussed. We found a "wavy" trend of green tide scales over the 14-year dissipation period. The dissipation direction was influenced by sea surface wind (SSW) and sea surface circulation (SSC). Under the action of northeast moving SSW and SSC, green tides move away from the shore and drift northward; under the action of northwest or southwest moving SSW and SSC, they moved towards the shore and drift southward. The date of dissipation was influenced by sea surface temperature (SST) and precipitation, and high SST accelerated the process of green tide dissipation, while precipitation slowed it down.

Remote sensing methods for biomass estimation of green algae attached to nursery-nets and raft rope.

Accurate estimation of the biomass of raft-attached green algae is important for predicting the scale of green-tides in the Yellow Sea, China. In this study, two different biomass estimation methods are proposed: green algae attached to nursery-net (GAAN) and green algae attached to rope (GAAR). The GAAN method involves the use of images obtained using an unmanned aerial vehicle (UAV), high-resolution satellite images, and data from a statistical yearbook. The GAAR method uses high-resolution satellite images and data from a field sample survey. The results showed that the biomass of GAAN and GAAR in the Subei Shoal during 2017 was 8868 tons and 2974 tons respectively. A longer-term study of the biomass of GAAN and GAAR could provide quantitative information for the earnings forecasts of Porphyra yezoensis and for green-tide prevention.

A UAV and S2A data-based estimation of the initial biomass of green algae in the South Yellow Sea.

Previous studies have shown that the initial biomass of green tide was the green algae attaching to Pyropia aquaculture rafts in the Southern Yellow Sea. In this study, the green algae was identified with unmanned aerial vehicle (UAV), an biomass estimation model was proposed for green algae biomass in the radial sand ridge area based on Sentinel-2A image (S2A) and UAV images. The result showed that the green algae was detected highly accurately with the normalized green-red difference index (NGRDI); approximately 1340 tons and 700 tons of green algae were attached to rafts and raft ropes respectively, and the lower biomass might be the main cause for the smaller scale of green tide in 2017. In addition, UAV play an important role in raft-attaching green algae monitoring and long-term research of its biomass would provide a scientific basis for the control and forecast of green tide in the Yellow Sea.

GIS-based health assessment of the marine ecosystem in Laizhou Bay, China.

According to 2014-2016 monitoring data, an assessment index system including water quality, depositional environment and ecosystem was built to evaluate the health statue of marine ecosystem in the Laizhou Bay using analytic hierarchy process (AHP) method. The results, spatialized in ArcGIS software, show: while the comprehensive ecological health index is 0.62, the ecological environmental quality in the Laizhou Bay is in a sub-healthy state; the unhealthy area is mainly concentrated in southwestern inshore region, and impacted by serious environmental problems, such as water eutrophication and heavy metal pollution; the northwestern and southeastern inshore regions are in a sub-healthy state, while the eastern inshore and northern areas are in the healthiest state. The land-based pollutants that discharge into the sea may be the leading factors that are causing ecological environment deterioration in the Laizhou Bay, and the reclamation work ongoing around the port has exacerbated the ecological risk.