The efficiency of monochromatic light to drive photosynthesis drops rapidly at wavelengths longer than 685nm. The photosynthetic efficiency of these longer wavelengths can be improved by adding shorter wavelength light, a phenomenon known as the Emerson enhancement effect. The reverse effect, the enhancement of photosynthesis under shorter wavelength light by longer wavelengths, however, has not been well studied and is often thought to be insignificant. We quantified the effect of adding far-red light (peak at 735nm) to red/blue or warm-white light on the photosynthetic efficiency of lettuce (Lactuca sativa). Adding far-red light immediately increased quantum yield of photosystem II (Φ) of lettuce by an average of 6.5 and 3.6% under red/blue and warm-white light, respectively. Similar or greater increases in Φ were observed after 20min of exposure to far-red light. This longer-term effect of far-red light on Φ was accompanied by a reduction in non-photochemical quenching of fluorescence (NPQ), indicating that far-red light reduced the dissipation of absorbed light as heat. The increase in Φ and complementary decrease in NPQ is presumably due to preferential excitation of photosystem I (PSI) by far-red light, which leads to faster re-oxidization of the plastoquinone pool. This facilitates reopening of PSII reaction centers, enabling them to use absorbed photons more efficiently. The increase in Φ by far-red light was associated with an increase in net photosynthesis (P). The stimulatory effect of far-red light increased asymptotically with increasing amounts of far-red. Overall, our results show that far-red light can increase the photosynthetic efficiency of shorter wavelength light that over-excites PSII.

Far-red photons (701-750 nm) are abundant in sunlight but are considered inactive for photosynthesis and are thus excluded from the definition of photosynthetically active radiation (PAR; 400-700 nm). Several recent studies have shown that far-red photons synergistically interact with shorter wavelength photons to increase leaf photochemical efficiency. The value of far-red photons in canopy photosynthesis has not been studied. Here, we report the effects of far-red photons on single leaf and canopy photosynthesis in 14 diverse crop species. Adding far-red photons (up to 40%) to a background of shorter wavelength photons caused an increase in canopy photosynthesis equal to adding 400-700 nm photons.Far-red alone minimally increased photosynthesis. This indicates that far-red photons are equally efficient at driving canopy photosynthesis when acting synergistically with traditionally defined photosynthetic photons. Measurements made using LEDs with peak wavelength of 711, 723, or 746 nm showed that the magnitude of the effect was less at longer wavelengths. The consistent response among diverse species indicates that the mechanism is common in higher plants. These results suggest that far-red photons (701-750 nm) should be included in the definition of PAR. K E Y W O R D SChl d and f, Emerson enhancement, far-red, photosynthetically active radiation, photosystems, whole-plant/canopy photosynthesis

Far-red photons regulate shade avoidance responses and can have powerful effects on plant morphology and radiation capture. Recent studies have shown that far-red photons (700 to 750 nm) efficiently drive photosynthesis when added to traditionally defined photosynthetic photons (400-700 nm). But the long-term effects of far-red photons on canopy quantum yield have not yet been determined. We grew lettuce in a four-chamber, steady-state canopy gas-exchange system to separately quantify canopy photon capture, quantum yield for CO 2 fixation, and carbon use efficiency. These measurements facilitate a mechanistic understanding of the effect of far-red photons on the components of plant growth. Daytime photosynthesis and night-time respiration of lettuce canopies were continuously monitored from seedling to harvest in five replicate studies. Plants were grown under a background of either red/blue or white light, each background with or without 15% (50 mmol m −2 s −1) of far-red photons substituting for photons between 400 and 700 nm. All four treatments contained 31.5% blue photons, and an equal total photon flux from 400 to 750 nm of 350 mmol m −2 s −1. Both treatments with far-red photons had higher canopy photon capture, increased daily carbon gain (net photosynthesis minus respiration at night), and 29 to 31% more biomass than control treatments. Canopy quantum yield was similar among treatments (0.057 ± 0.002 mol of CO 2 fixed in gross photosynthesis per mole of absorbed photons integrated over 400 to 750 nm). Carbon use efficiency (daily carbon gain/gross photosynthesis) was also similar for mature plants (0.61 ± 0.02). Photosynthesis increased linearly with increasing photon capture and had a common slope among all four treatments, which demonstrates that the faster growth with far-red photon substitution was caused by enhanced photon capture through increased leaf expansion. The equivalent canopy quantum yield among

A crewed mission to and from Mars may include an exciting array of enabling biotechnologies that leverage inherent mass, power, and volume advantages over traditional abiotic approaches. In this perspective, we articulate the scientific and engineering goals and constraints, along with example systems, that guide the design of a surface biomanufactory. Extending past arguments for exploiting stand-alone elements of biology, we argue for an integrated biomanufacturing plant replete with modules for microbial in situ resource utilization, production, and recycling of food, pharmaceuticals, and biomaterials required for sustaining future intrepid astronauts. We also discuss aspirational technology trends in each of these target areas in the context of human and robotic exploration missions.

Linear electron transport depends on balanced excitation of photosystem I and II. Far-red light preferentially excites photosystem I (PSI) and can enhance the photosynthetic efficiency when combined with light that over-excites photosystem II (PSII). The efficiency of different wavelengths of far-red light exciting PSI was quantified by measuring the change in quantum yield of PSII (Φ ) of lettuce (Lactuca sativa) under red/blue light with narrowband far-red light added (from 678 to 752 nm, obtained using laser diodes). The Φ of lettuce increased with increasing wavelengths of added light from 678 to 703 nm, indicating longer wavelengths within this region are increasingly used more efficiently by PSI than by PSII. Adding 721 nm light resulted in similar Φ as adding 703 nm light, but Φ tended to decrease as wavelength increased from 721 to 731 nm, likely due to decreasing absorptance and low photon energy. Adding 752 nm light did not affect Φ . Leaf chlorophyll fluorescence light response measurements showed lettuce had higher Φ under halogen light (rich in far-red) than under red/blue light (which over-excites PSII). Far-red light is more photosynthetically active than commonly believed, because of its synergistic interaction with light of shorter wavelengths.