High nitrogen inhibits biomass and saponins accumulation in a medicinal plant Panax notoginseng.

Nitrogen (N) is an important macronutrient and is comprehensively involved in the synthesis of secondary metabolites. However, the interaction between N supply and crop yield and the accumulation of effective constituents in an N-sensitive medicinal plant Panax notoginseng (Burkill) F. H. Chen is not completely known. Morphological traits, N use and allocation, photosynthetic capacity and saponins accumulation were evaluated in two- and three-year-old P. notoginseng grown under different N regimes. The number and length of fibrous root, total root length and root volume were reduced with the increase of N supply. The accumulation of leaf and stem biomass (above-ground) were enhanced with increasing N supply, and LN-grown plants had the lowest root biomass. Above-ground biomass was closely correlated with N content, and the relationship between root biomass and N content was negatives in P. notoginseng (r = -0.92). N use efficiency-related parameters, NUE (N use efficiency, etc.), NC (N content in carboxylation system component) and P n (the net photosynthetic rate) were reduced in HN-grown P. notoginseng. SLN (specific leaf N), Chl (chlorophyll), NL (N content in light capture component) increased with an increase in N application. Interestingly, root biomass was positively correlated with NUE, yield and P n. Above-ground biomass was close negatively correlated with photosynthetic N use efficiency (PNUE). Saponins content was positively correlated with NUE and P n. Additionally, HN improved the root yield of per plant compared with LN, but reduced the accumulation of saponins, and the lowest yield of saponins per unit area (35.71 kg·hm-2) was recorded in HN-grown plants. HN-grown medicinal plants could inhibit the accumulation of root biomass by reducing N use and photosynthetic capacity, and HN-induced decrease in the accumulation of saponins (C-containing metabolites) might be closely related to the decline in N efficiency and photosynthetic capacity. Overall, N excess reduces the yield of root and C-containing secondary metabolites (active ingredient) in N-sensitive medicinal species such as P. notoginseng.

A Comparison of Photoprotective Mechanism in Different Light-Demanding Plants Under Dynamic Light Conditions.

Light intensity is highly heterogeneous in nature, and plants have evolved a series of strategies to acclimate to dynamic light due to their immobile lifestyles. However, it is still unknown whether there are differences in photoprotective mechanisms among different light-demanding plants in response to dynamic light, and thus the role of non-photochemical quenching (NPQ), electron transport, and light energy allocation of photosystems in photoprotection needs to be further understood in different light-demanding plants. The activities of photosystem II (PSII) and photosystem I (PSI) in shade-tolerant species Panax notoginseng, intermediate species Polygonatum kingianum, and sun-demanding species Erigeron breviscapus were comparatively measured to elucidate photoprotection mechanisms in different light-demanding plants under dynamic light. The results showed that the NPQ and PSII maximum efficiency (F v'/F m') of E. breviscapus were higher than the other two species under dynamic high light. Meanwhile, cyclic electron flow (CEF) of sun plants is larger under transient high light conditions since the slope of post-illumination, P700 dark reduction rate, and plastoquinone (PQ) pool were greater. NPQ was more active and CEF was initiated more readily in shade plants than the two other species under transient light. Moreover, sun plants processed higher quantum yield of PSII photochemistry (ΦPSII), quantum yield of photochemical energy conversion [Y(I)], and quantum yield of non-photochemical energy dissipation due to acceptor side limitation (Y(NA), while the constitutive thermal dissipation and fluorescence (Φf,d) and quantum yield of non-photochemical energy dissipation due to donor side limitation [Y(ND)] of PSI were higher in shade plants. These results suggest that sun plants had higher NPQ and CEF for photoprotection under transient high light and mainly allocated light energy through ΦPSII and ΦNPQ, while shade plants had a higher Φf,d and a larger heat dissipation efficiency of PSI donor. Overall, it has been demonstrated that the photochemical efficiency and photoprotective capacity are greater in sun plants under transient dynamic light, while shade plants are more sensitive to transient dynamic light.

Responses of Linear and Cyclic Electron Flow to Nitrogen Stress in an N-Sensitive Species Panax notoginseng.

Nitrogen (N) is a primary factor limiting leaf photosynthesis. However, the mechanism of N-stress-driven photoinhibition of the photosystem I (PSI) and photosystem II (PSII) is still unclear in the N-sensitive species such as Panax notoginseng, and thus the role of electron transport in PSII and PSI photoinhibition needs to be further understood. We comparatively analyzed photosystem activity, photosynthetic rate, excitation energy distribution, electron transport, OJIP kinetic curve, P700 dark reduction, and antioxidant enzyme activities in low N (LN), moderate N (MN), and high N (HN) leaves treated with linear electron flow (LEF) inhibitor [3-(3,4-dichlorophenyl)-1,1-dimethyl urea (DCMU)] and cyclic electron flow (CEF) inhibitor (methyl viologen, MV). The results showed that the increased application of N fertilizer significantly enhance leaf N contents and specific leaf N (SLN). Net photosynthetic rate (P n) was lower in HN and LN plants than in MN ones. Maximum photochemistry efficiency of PSII (F v/F m), maximum photo-oxidation P700+ (P m), electron transport rate of PSI (ETRI), electron transport rate of PSII (ETRII), and plastoquinone (PQ) pool size were lower in the LN plants. More importantly, K phase and CEF were higher in the LN plants. Additionally, there was not a significant difference in the activity of antioxidant enzyme between the MV- and H2O-treated plants. The results obtained suggest that the lower LEF leads to the hindrance of the formation of ΔpH and ATP in LN plants, thereby damaging the donor side of the PSII oxygen-evolving complex (OEC). The over-reduction of PSI acceptor side is the main cause of PSI photoinhibition under LN condition. Higher CEF and antioxidant enzyme activity not only protected PSI from photodamage but also slowed down the damage rate of PSII in P. notoginseng grown under LN.

Responses of photosystem to long-term light stress in a typically shade-tolerant species Panax notoginseng.

Photosynthetic adaptive strategies vary with the growth irradiance. The potential photosynthetic adaptive strategies of shade-tolerant species Panax notoginseng (Burkill) F. H. Chen to long-term high light and low light remains unclear. Photosynthetic performance, photosynthesis-related pigments, leaves anatomical characteristics and antioxidant enzyme activities were comparatively determined in P. notoginseng grown under different light regimes. The thickness of the upper epidermis, palisade tissue, and lower epidermis were declined with increasing growth irradiance. Low-light-grown leaves were declined in transpiration rate (Tr) and stomatal conductance (Cond), but intercellular CO2 concentration (C i) and net photosynthesis rate (P n) had opposite trends. The maximum photo-oxidation P 700 + (P m) was greatly reduced in 29.8% full sunlight (FL) plants; The maximum quantum yield of photosystem II (F v/F m) in 0.2% FL plants was significantly lowest. Electron transport, thermal dissipation, and the effective quantum yield of PSI [Y(I)] and PSII [Y(II)] were declined in low-light-grown plants compared with high-light-grown P. notoginseng. The minimum value of non-regulated energy dissipation of PSII [Y(NO)] was recorded in 0.2% FL P. notoginseng. OJIP kinetic curve showed that relative variable fluorescence at J-phase (V J) and the ratio of variable fluorescent F K occupying the F J-F O amplitude (W k) were significantly increased in 0.2% FL plants. However, the increase in W k was lower than the increase in V J. In conclusion, PSI photoinhibition is the underlying sensitivity of the typically shade-tolerant species P. notoginseng to high light, and the photodamage to PSII acceptor side might cause the typically shade-tolerant plants to be unsuitable for long-term low light stress.

The Responses of Light Reaction of Photosynthesis to Dynamic Sunflecks in a Typically Shade-Tolerant Species Panax notoginseng.

Light is highly heterogeneous in natural conditions, and plants need to evolve a series of strategies to acclimate the dynamic light since it is immobile. The present study aimed to elucidate the response of light reaction of photosynthesis to dynamic sunflecks in a shade-tolerant species Panax notoginseng and to examine the regulatory mechanisms involved in an adaptation to the simulated sunflecks. When P. notoginseng was exposed to the simulated sunflecks, non-photochemical quenching (NPQ) increased rapidly to the maximum value. Moreover, in response to the simulated sunflecks, there was a rapid increase in light-dependent heat dissipation quantum efficiency of photosystem II (PSII) (ΦNPQ), while the maximum quantum yield of PSII under light (F v'/F m') declined. The relatively high fluorescence and constitutive heat dissipation quantum efficiency of PSII (Φf,d) in the plants exposed to transient high light (400, 800, and 1,600 µmol m-2 s-1) was accompanied by the low effective photochemical quantum yield of PSII (ΦPSII) after the dark recovery for 15 min, whereas the plants exposed to transient low light (50 µmol m-2 s-1) has been shown to lead to significant elevation in ΦPSII after darkness recovery. Furthermore, PSII fluorescence and constitutive heat dissipation electron transfer rate (J f,d) was increased with the intensity of the simulated sunflecks, the residual absorbed energy used for the non-net carboxylative processes (J NC) was decreased when the response of electron transfer rate of NPQ pathway of PSII (J NPQ) to transient low light is restricted. In addition, the acceptor-side limitation of PSI [Y(NA)] was increased, while the donor-side limitation of photosystems I (PSI) [Y(ND)] was decreased at transient high light conditions accompanied with active cyclic electron flow (CEF). Meanwhile, when the leaves were exposed to transient high light, the xanthophyll cycle (V cycle) was activated and subsequently, the J NPQ began to increase. The de-epoxidation state [(Z + A)/(V + A + Z)] was strongly correlated with NPQ in response to the sunflecks. In the present study, a rapid engagement of lutein epoxide (Lx) after the low intensity of sunfleck together with the lower NPQ contributed to an elevation in the maximum photochemical quantum efficiency of PSII under the light. The analysis based on the correlation between the CEF and electron flow devoted to Ribulose-1, 5-bisphosphate (RuBP) oxygenation (J O) indicated that at a high light intensity of sunflecks, the electron flow largely devoted to RuBP oxygenation would contribute to the operation of the CEF. Overall, photorespiration plays an important role in regulating the CEF of the shade-tolerant species, such as P. notoginseng in response to transient high light, whereas active Lx cycle together with the decelerated NPQ may be an effective mechanism of elevating the maximum photochemical quantum efficiency of PSII under light exposure to transient low light.