The role of soil in the contribution of food and feed.

Soils play a critical role in the production of food and feed for a growing global population. Here, we review global patterns in soil characteristics, agricultural production and the fate of embedded soil nutrients. Nitrogen- and organic-rich soils supported the highest crop yields, yet the efficiency of nutrient utilization was concentrated in regions with lower crop productivity and lower rates of chemical fertilizer inputs. Globally, soil resources were concentrated in animal feed, resulting in large inefficiencies in nutrient utilization and losses from the food system. Intercontinental transport of soil-derived nutrients displaced millions of tonnes of nitrogen and phosphorus annually, much of which was ultimately concentrated in urban waste streams. Approximately 40% of the global agricultural land area was in small farms providing over 50% of the world's food and feed needs but yield gaps and economic constraints limit the ability to intensify production on these lands. To better use and protect soil resources in the global food system, policies and actions should encourage shifts to more nutrient-efficient diets, strategic intensification and technological improvement, restoration and maintenance of soil fertility and stability, and enhanced resilience in the face of global change. This article is part of the theme issue 'The role of soils in delivering Nature's Contributions to People'.

Soil organic carbon is not just for soil scientists: measurement recommendations for diverse practitioners.

Soil organic carbon (SOC) regulates terrestrial ecosystem functioning, provides diverse energy sources for soil microorganisms, governs soil structure, and regulates the availability of organically bound nutrients. Investigators in increasingly diverse disciplines recognize how quantifying SOC attributes can provide insight about ecological states and processes. Today, multiple research networks collect and provide SOC data, and robust, new technologies are available for managing, sharing, and analyzing large data sets. We advocate that the scientific community capitalize on these developments to augment SOC data sets via standardized protocols. We describe why such efforts are important and the breadth of disciplines for which it will be helpful, and outline a tiered approach for standardized sampling of SOC and ancillary variables that ranges from simple to more complex. We target scientists ranging from those with little to no background in soil science to those with more soil-related expertise, and offer examples of the ways in which the resulting data can be organized, shared, and discoverable.

Long-term patterns in tropical reforestation: plant community composition and aboveground biomass accumulation.

Primary tropical forests are renowned for their high biodiversity and carbon storage, and considerable research has documented both species and carbon losses with deforestation and agricultural land uses. Economic drivers are now leading to the abandonment of agricultural lands, and the area in secondary forests is increasing. We know little about how long it takes for these ecosystems to achieve the structural and compositional characteristics of primary forests. In this study, we examine changes in plant species composition and aboveground biomass during eight decades of tropical secondary succession in Puerto Rico, and compare these patterns with primary forests. Using a well-replicated chronosequence approach, we sampled primary forests and secondary forests established 10, 20, 30, 60, and 80 years ago on abandoned pastures. Tree species composition in all secondary forests was different from that of primary forests and could be divided into early (10-, 20-, and 30-year) vs. late (60- and 80-year) successional phases. The highest rates of aboveground biomass accumulation occurred in the first 20 years, with rates of C sequestration peaking at 6.7 +/- 0.5 Mg C x ha(-1) x yr(-1). Reforestation of pastures resulted in an accumulation of 125 Mg C/ha in aboveground standing live biomass over 80 years. The 80 year-old secondary forests had greater biomass than the primary forests, due to the replacement of woody species by palms in the primary forests. Our results show that these new ecosystems have different species composition, but similar species richness, and significant potential for carbon sequestration, compared to remnant primary forests.

Evidence for nicotinic acetylcholine receptors on nasal trigeminal nerve endings of the rat.

The peripheral chemoreceptors of the trigeminal system in the nasal cavity are presumed to be free nerve endings arising from Adelta and C fibers. These fibers appear to be scattered throughout the nasal epithelium, and arise from the nasopalatine and ethmoid branches of the trigeminal nerve. In the present study, the effects of nicotinic acetylcholine receptor (nAChR) blockers on ethmoid nerve responses to nicotine and cyclohexanone were examined. Multiunit neural recordings were obtained from the ethmoid nerve of Sprague-Dawley rats. Vapor-phase nicotine (12.5 p.p.m.) and cyclohexanone (450 p.p. m.) were delivered to the rats' nares via an air-dilution olfactometer. The magnitude of the response to nicotine decreased after the administration of the nAChR blockers dihydro-beta-erythroidine hydrobromide (DHBE) and mecamylamine hydrochloride. DHBE is a competitive nicotinic receptor antagonist specific for the alpha4beta2 receptor subtype and mecamylamine is known to bind alpha3beta4 and alpha4beta2 receptors. The nAChR blockers had no effect on ethmoid nerve responses to cyclohexanone. These results suggest that the mechanism by which at least one irritant stimulates nasal trigeminal nerve endings involves the binding of irritant with a specific receptor.

Odorants presented to the rat nasal cavity increase cortical blood flow.

Complaints about unpleasant environmental odorants, both outdoor and indoor, are increasingly being reported. The main complaints of health symptoms from environmental odorants are eye, nose and throat irritation, headache and drowsiness. Complaints may arise from the stimulation of olfactory receptors or trigeminal chemoreceptors. Stimulation of cerebrovascular nociceptors originating from a branch of the trigeminal nerve may be associated with an increase in cortical blood flow which is thought to be related to headache. Since odorants are reported to elicit headaches, the possibility that odorants may increase cortical blood flow was examined. Cortical blood flow was monitored in rats using a laser-Doppler flowmeter. The flowmeter probe was placed over the left frontal cortex while propionic acid, cyclohexanone, amyl acetate or butanol was delivered to the nasal cavity via an olfactometer. Cortical blood flow increased as the concentration increased for three of the odorants tested. The greatest increase in blood flow occurred to the presentation of propionic acid, followed by cyclohexanone and amyl acetate. There was no response to butanol. These data demonstrate that odorants can alter cerebrovascular blood flow, which may account, in part, for one of the health symptoms reported for odorants.

The effects of neonatal capsaicin administration on trigeminal nerve chemoreceptors in the rat nasal cavity.

Trigeminal nerve fibers in the nasal cavity respond to a variety of volatile chemical stimuli. Some of these trigeminal nerve fibers have been suggested to be capsaicin-sensitive and thus belong to a class of pain receptor rather than constituting a separate class of chemoreceptor. Our current results confirm this suggestion. Trigeminal nerve responses to volatile chemical stimuli were eliminated in rats which were injected with capsaicin on the second day of life. Animals whose nerves were unresponsive to chemical stimuli also exhibited a loss of intraepithelial peptide-immunoreactive fibers in their nasal cavities. The results of this study suggest that trigeminal nerve fibers in the nasal cavity which respond to chemical stimuli may be polymodal nociceptors which contain substance P, calcitonin gene-related peptide, or perhaps other neuropeptides.

Ultrastructure of substance P- and CGRP-immunoreactive nerve fibers in the nasal epithelium of rodents.

The respiratory and olfactory mucosae of rats and mice were examined at ultrastructural levels for the presence of intraepithelial nerve endings. Immunocytochemical studies utilizing antisera directed against substance P and calcitonin gene-related peptide (CGRP) revealed numerous intraepithelial peptide-immunoreactive fibers near the basal region of the epithelium. Occasional transepithelial fibers were observed to extend outward to nearly reach the epithelial surface. In no cases, however, did the transepithelial fibers reach the surface, but instead, stopped at the line of tight junctions approximately 1 micron from the surface. No specialized contacts between the nerve fibers and the epithelial cells were observed. The transepithelial fibers provide a possible anatomical substrate for the sensitivity of the trigeminal nerve to many air-borne chemical stimuli. That potential chemical stimuli must traverse the tight-junctional barrier may explain why lipid solubility is related to effectiveness for trigeminal stimuli.

A comparison of the discriminatory ability and sensitivity of the trigeminal and olfactory systems to chemical stimuli in the tiger salamander.

Trigeminal receptors can respond to a wide variety of chemical stimuli, but it is unknown whether these receptors mediate discrimination between chemical stimuli matched for equal perceptual intensity. The present electrophysiological and behavioral experiments address this issue using tiger salamanders, Ambystoma tigrinum, and four compounds (amyl acetate, cyclohexanone, butanol, and d-limonene). In addition, the relative sensitivities of the trigeminal and olfactory systems to these compounds are compared. In electrophysiological cross-adaptation experiments (amyl acetate vs cyclohexanone; butanol vs d-limonene), there was complete cross adaptation such that only concentrations above the background (cross-adapting) stimulus concentration elicited responses, suggesting that chemical stimuli may stimulate trigeminal receptors nonspecifically. In behavioral experiments (amyl acetate vs cyclohexanone; butanol vs d-limonene), only animals with intact olfactory nerves could discriminate between perceptually equivalent concentrations, that is concentrations that elicited the same level of responding. Both electrophysiologically and behaviorally, the trigeminal system exhibited higher thresholds than the olfactory system. We conclude that trigeminal chemoreceptors, at least in salamanders, are unable to discriminate between these two pairs of compounds when matched for equal perceptual intensity, and that trigeminal chemoreceptors are less sensitive than olfactory receptors.

Capsaicin and its effects on olfaction and trigeminal chemoreception.

Capsaicin injections severely reduced or eliminated nasal trigeminal responses to 3 odorants (Experiment 1). However, capsaicin treated animals exhibited no deficits in locating buried food, in odor avoidance learning, or in operant odor detection and discrimination (Experiments 2 and 3). In addition, capsaicin desensitization did not affect responsiveness to salty or sour, but may have raised rejection thresholds for bitter (Experiment 4). Finally, while desensitized animals rejected menthol solution, they consumed relatively more than controls, suggesting that capsaicin may have menthol sensitivity. The present results suggest that substance P-containing fibers mediate trigeminal responsiveness to odorants and irritants but that the loss of this responsiveness does not appreciably affect smell or taste, per se.

Nasal trigeminal chemoreception: responses to n-aliphatic alcohols.

Odorant molecules can stimulate nasal trigeminal receptors, but the properties of such molecules which make them effective stimuli are largely unknown. In the present study, we obtained integrated multiunit responses from the ethmoid branch of the rat trigeminal nerve to a homologous series of aliphatic alcohols. Our aim was to determine whether lipid solubility might correlate with stimulus efficacy. Response thresholds (ranging from 3000 ppm for methanol to 3 ppm for octanol) decreased with increasing carbon chain length, suggesting that lipid solubility is important for stimulus effectiveness. One plausible explanation for the importance of lipophilicity is that the more lipid soluble a substance, the more easily it can penetrate epithelial layers to reach chemoreceptive trigeminal nerve endings. Since all stimuli at vapor saturation elicited responses within 0.5 s, and because diffusion of stimulus molecules through epithelium is slow, we speculate that trigeminal nerve endings lie closer to the epithelial surface than previously thought.

Olfactory responses of aquatic and terrestrial tiger salamanders to airborne and waterborne stimuli.

Electro-olfactograms (EOGs) were used to assess olfactory responding by aquatic larval and terrestrial adult tiger salamanders (Ambystoma tigrinum) to airborne volatile compounds, and volatile and non-volatile compounds in aqueous solution. Both forms of salamander showed saturation effects to presentations of airborne stimuli (Fig. 2). Saturation was not observed, however, to stimulus presentations in aqueous solution (Figs. 2, 3). When threshold values and concentration-response curve parameters were compared, non-volatile amino acids in solution were more potent stimuli for larvae while airborne volatiles were more potent stimuli for adults (Tables 1, 2). We infer that metamorphosis in the tiger salamander is accompanied by changes in olfactory response characteristics, due possibly to changes in receptor population, changes in perireceptor properties (e.g. mucus) or to changes in stimulus access.

Inhibition by amiloride of chorda tympani responses evoked by monovalent salts.

The diuretic, amiloride, is a potent yet reversible inhibitor of passive sodium transport in many epithelia. It has been shown to inhibit sodium transport in dorsal lingual epithelia and to inhibit both psychophysical and neural measures of salt taste. The present results demonstrate that amiloride's action as an inhibitor of integrated whole chorda tympani nerve recordings in the rat is specific for Li and Na salts, displaying little inhibition of neural responses evoked by KCl and RbCl. Amiloride reduces both the phasic and tonic portion of the nerve recording equally. When amiloride inactivates the tonic portion of the nerve response to 250 mM NaCl, only a portion of the response is affected. Complete inactivation does not occur even at high amiloride concentrations. With amiloride flowing constantly over the tongue at 1 microM, 10 microM, or 50 microM a reciprocal plot of stimulus NaCl concentration versus response is non-linear. This result suggests that the inhibition of the NaCl-induced response has both competitive and non-competitive properties. These results support the hypothesis that salt taste is mediated in part by amiloride sensitive Na-channels located in taste receptor cell plasma membranes.

Rat trigeminal, olfactory and taste responses after capsaicin desensitization.

Experiment 1 showed that capsaicin injections severely reduced or eliminated nasal trigeminal responses to 3 odorants. Experiments 2 and 3 investigated whether desensitized animals could behaviorally detect and discriminate odors. Capsaicin treated animals had no measurable deficits in locating buried food, in odor aversion learning, or in operant odor detection and discrimination. Experiment 4 examined whether behavioral responsiveness to salty, sour and bitter tastes was affected by desensitization. Capsaicin injections did not affect responsiveness to salty or sour, but may have raised rejection thresholds for bitter. Broadly, the present results suggest that substance P-containing fibers mediate trigeminal responsiveness to odorants and irritants but that the loss of this responsiveness does not appreciably affect smell or taste, per se.

Trigeminally mediated odor aversions in starlings.

Electrophysiological and psychophysical evidence has demonstrated that trigeminal receptors in the nasal cavity respond to odorants. Despite these demonstrations of trigeminal chemoreception, it is not clear whether naso-trigeminal stimulation can be used to mediate learned behaviors in contexts such as feeding. Here, we report that starlings will learn aversions to odorant volatiles in a feeding context and that these aversions can be mediated by trigeminal cues.

Chemosensitivity of rat nasal trigeminal receptors.

Electrophysiological responses to odorants delivered via an air dilution olfactometer were recorded from the ethmoid branch of the trigeminal nerve innervating the nasal cavity. Thresholds were obtained for nine compounds with those for heptanol (21-137 ppm) and propionic acid (39-49) ppm consistently being the lowest. Not all odorants e.g., phenethyl alcohol, elicited responses in all rats even at vapor saturation. A striking degree of correlation was present between the rat whole-nerve electrophysiological response magnitudes of this study and the human anosmic intensity ratings established in the work of Doty et al. [9] to vapor saturated stimuli. These results suggest that the rat is an excellent model for assessing the stimulatory effectiveness of odorants on human trigeminal receptors. The possible role of the trigeminal system in the perception of odors as well as the physiologic effects of odorants due to trigeminal stimulation are discussed.

The underwater electro-olfactogram: a tool for the study of the sense of smell of marine fishes.

Recording the olfactory receptor activity of marine fishes presents problems due to the shunting of the electrical signals by the highly conductive sea water, which results in significant signal loss. By recording the large signal-to-noise ratio D. C. potentials using the underwater electro-olfactogram (EOG), we were able to study olfactory receptor properties of freshwater and marine fishes in a comparable manner.