Fertile forests produce biomass more efficiently.

Trees with sufficient nutrition are known to allocate carbon preferentially to aboveground plant parts. Our global study of 49 forests revealed an even more fundamental carbon allocation response to nutrient availability: forests with high-nutrient availability use 58 ± 3% (mean ± SE; 17 forests) of their photosynthates for plant biomass production (BP), while forests with low-nutrient availability only convert 42 ± 2% (mean ± SE; 19 forests) of annual photosynthates to biomass. This nutrient effect largely overshadows previously observed differences in carbon allocation patterns among climate zones, forest types and age classes. If forests with low-nutrient availability use 16 ± 4% less of their photosynthates for plant growth, what are these used for? Current knowledge suggests that lower BP per unit photosynthesis in forests with low- versus forests with high-nutrient availability reflects not merely an increase in plant respiration, but likely results from reduced carbon allocation to unaccounted components of net primary production, particularly root symbionts.

Light and nitrogen competition limit Lolium perenne in experimental grasslands of increasing plant diversity.

Positive species richness effects on aboveground community productivity in experimental grasslands have been reported to correlate with variable responses of individual species. So far, it is largely unknown whether more complete use of resources at the community level correlates with resource limitation of particular species and may explain their decreasing performance with increasing plant diversity. Using the subordinate grass species Lolium perenne L. as a model, we monitored populations in 82 experimental grasslands of different plant diversity (Jena Experiment) from year 2 to 6 after establishment, and measured ecophysiological leaf traits related to light and nutrient acquisition and use. Population and plant individual sizes of L. perenne decreased with increasing species richness. A decrease in transmitted light with increasing species richness and legume proportion correlated with increasing specific leaf area (SLA). Despite this morphological adaptation to lower light availability, decreasing foliar δ(13) C signatures with increasing species richness and low variation in leaf gas exchange and chlorophyll concentrations suggested a low capacity of L. perenne for adjustment to canopy shade. Leaf nitrogen concentrations and foliar δ(15) N signatures indicated a better N supply in communities with legumes and a shift in the uptake of different N forms with increasing species richness. Leaf blade nitrate and carbohydrate concentrations as indicators of plants nutritional status supported that light limitation with increasing species richness and legume proportions, combined with a N limitation in communities with increasing proportions of non-legumes, correlated with the decreasing performance of L. perenne in communities of increasing plant diversity.

The influence of climate and fructification on the inter-annual variability of stem growth and net primary productivity in an old-growth, mixed beech forest.

The periodic production of large seed crops by trees (masting) and its interaction with stem growth has long been the objective of tree physiology research. However, very little is known about the effects of masting on stem growth and total net primary productivity (NPP) at the stand scale. This study was conducted in an old-growth, mixed deciduous forest dominated by Fagus sylvatica (L.) and covers the period from 2003 to 2007, which comprised wet, dry and regular years as well as two masts of Fagus and one mast of the co-dominant tree species Fraxinus excelsior (L.) and Acer pseudoplatanus (L.). We combined analyses of weather conditions and stem growth at the tree level (inter- and intra-annual) with fruit, stem and leaf production, and estimates of total NPP at the stand level. Finally, we compared the annual demand of carbon for biomass production with net canopy assimilation (NCA), derived from eddy covariance flux measurements, chamber measurements and modelling. Annual stem growth of Fagus was most favoured by warm periods in spring and that of Fraxinus by high precipitation in June. For stem growth of Acer and for fruit production, no significant relationships with mean weather conditions were found. Intra-annual stem growth of all species was strongly reduced when the relative plant-available water in soil dropped below a threshold of about 60% between May and July. The inter-annual variations of NCA, total NPP and leaf NPP at the stand level were low (mean values 1313, 662 and 168 g C m(-2) year(-1), respectively), while wood and fruit production varied more and contrarily (wood: 169-241 g C m(-2) year(-1); fruits: 21-142 g C m(-2) year(-1)). In all years, an annual surplus of newly assimilated carbon was calculated (on average 100 g C m(-2) year(-1)). The results suggest that stem growth is generally not limited by insufficient carbon resources; only in mast years a short-term carbon shortage may occur in spring. In contrast to common assumption, stem growth alone is not a sufficient proxy for total biomass production or the control of carbon sequestration by weather extremes.

Neurophysiological studies of flight-related density-dependent phase characteristics in locusts.

Locusts show an extreme example of density-dependent phase polymorphism, demonstrating within the species differences in morphology as well as biology, dependent on the population density. Behavior is the primary density-dependent change which facilitates the appearance of various morphological and physiological phase characteristics. We have studied density dependent differences in flight related sensory and central neural elements in the desert locust Schistocerca gregaria. Wind generated high frequency spiking activity in the tritocerebral commissure giant (TCG, an identified interneuron that relay inputs from head hair receptors to thoracic motor centers) that was much less intense in solitary locusts, compared to gregarious ones. In addition the solitary locusts' TCG demonstrated much stronger adaptation of its response. In cases when flight was initiated high frequency TCG activity was independent of the locust phase. The tritocerebral commissure dwarf (TCD) is a GABAergic flight related interneuron that is sensitive to ambient illumination intensity. An increase in the TCD spontaneous activity under dark vs. light conditions was significantly higher in gregarious locusts then in solitary ones, implying a flight-related inhibitory mechanism that is far more active in gregarious locusts under dark conditions. Thus, density-dependent phase differences in interneuron activity pattern and properties well reflect and may be at least partially responsible to behavioral flight-related characteristics.

Auditory-evoked evasive manoeuvres in free-flying locusts and moths.

We presented free-flying locusts (Locusta migratoria L.) with sounds that varied in temporal structure and carrier frequency as they flew toward a light source in a flight room under controlled temperature and light conditions. Previous studies have shown tethered locusts react more often to trains of 30-kHz pulses than to pulse trains below 10 kHz. Further, this acoustic startle response has been suggested to function in bat-avoidance. We expected free-flying locusts to respond similarly; however, we found locusts responded to all sounds we presented, not just high-frequency, "bat-like" sounds. Response rates of turns, loops, and dives varied from 6% to 26% but were statistically independent of carrier frequency and/or pulse structure. Free-flying moths and tethered locusts were tested using a subset of our acoustic stimuli under the same temperature and light conditions as the free-flying locusts. Moth responses were carrier frequency dependent as were responses of tethered locusts positioned along the flight path observed in our free-flight trials. All responses were unaffected by a 90% reduction in room light. We conclude that locusts possess an acoustic startle response evocable in free flight, however, free-flying locusts do not show the same discrimination observed in tethered locusts under similar conditions.

Neural correlates to flight-related density-dependent phase characteristics in locusts.

Locust phase polymorphism is an extreme example of behavioral plasticity; in response to changes in population density, locusts dramatically alter their behavior. These changes in behavior facilitate the appearance of various morphological and physiological phase characteristics. One of the principal behavioral changes is the more intense flight behavior and improved flight performance of gregarious locusts compared to solitary ones. Surprisingly, the neurophysiological basis of the behavioral phase characteristics has received little attention. Here we present density-dependent differences in flight-related sensory and central neural elements in the desert locust. Using techniques already established for gregarious locusts, we compared the response of locusts of both phases to controlled wind stimuli. Gregarious locusts demonstrated a lower threshold for wind-induced flight initiation. Wind-induced spiking activity in the locust tritocerebral commissure giants (TCG, a pair of identified interneurons that relay input from head hair receptors to thoracic motor centers) was found to be weaker in solitary locusts compared to gregarious ones. The solitary locusts' TCG also demonstrated much stronger spike frequency adaptation in response to wind stimuli. Although the number of forehead wind sensitive hairs was found to be larger in solitary locusts, the stimuli conveyed to their flight motor centers were weaker. The tritocerebral commissure dwarf (TCD) is an inhibitory flight-related interneuron in the locust that responds to light stimuli. An increase in TCD spontaneous activity in dark conditions was significantly stronger in gregarious locusts than in solitary ones. Thus, phase-dependent differences in the activity of flight-related interneurons reflect behavioral phase characteristics.

Relationships between body mass, motor output and flight variables during free flight of juvenile and mature adult locusts, Schistocerca gregaria.

Little information is available about how the adult locust flight system manages to match the aerodynamic demands that result from an increase in body mass during postmoult maturation. In Schistocerca gregaria of both sexes, flight variables, including flight speed, ascent angle and body angle, were investigated under closed-loop conditions (i.e. during free flight) as a function of adult maturation. Motor patterns were examined by telemetric electromyography in juvenile and adult mature animals of both sexes. Functional relationships between particular flight variables were investigated by additional loading of the animals and by reductions in wing area. The results indicate that an increase in flight speed as the flight system matures enables it to match the aerodynamic demands resulting from increases in body mass. Furthermore, the data suggest that this postmoult increase in flight speed is not simply a consequence of the increase in wingbeat frequency observed during maturation. The instantaneous body angle during flight is controlled mainly by aerodynamic output from the wings. In addition, the mean body angle decreases during maturation in both sexes, and this may play an important part in the directional control of the resultant flight force vector.

Timing of elevator muscle activity during climbing in free locust flight

Despite detailed knowledge of the sensory-motor interactions during elevator muscle timing for the generation of a 'functional' flight motor pattern in flying locusts, there is little information about how a possible shift in the onset of elevator activity is correlated with changes in flight variables under closed-loop conditions (i.e. during free flight). Free-flight variables were investigated with respect to ascent angle during climbing flight in locusts Schistocerca gregaria. The motor pattern during free flight was examined by telemetric electromyography of particular antagonistic flight muscles in both ipsilateral hemisegments of the pterothorax while flight variables were recorded simultaneously on video. In the majority of the animals tested, the onset of elevator muscle activity within the wingbeat cycle is delayed when animals increase their ascent angle during climbing flight. In accordance with the motor pattern, the downstroke phase and the stroke amplitude of the wings increased with increasing the ascent angle. This suggests that the relative elevator timing during the wingbeat cycle may be related to the generation of the additional aerodynamic lift required for ascending flight and may, therefore, play a role in the regulation of ascent angle during free flight in the locust.

A radiotelemetric 2-channel unit for transmission of muscle potentials during free flight of the desert locust, Schistocerca gregaria.

A new radio-telemetric technique for neuroethological investigation of the natural behaviour of insects is presented. The 2-channel miniature transmitter device allows the transmission of electromyograms of 2 muscles during free flight of a locust. The mass of this transmitter and power supply is 0.55 g and can be carried by a mature female without marked impairment of the free-flight behaviour. The radiated power of the transmitter is approximately 20 nW and is sufficient to cover a range of more than 20 m. The carrier-frequency (145 MHz) of the system is frequency-modulated by the inputs of the 2 different channels. Channel separation is achieved by a multivibrator circuit. The chopping frequency (2 kHz) allows a convenient resolution of both signal channels. The design of the 2-channel transmitter device is presented and tested. Its relevance for the studies of natural flight of locusts is given, however, other applications are also feasible.

Segmental differentiation processes in embryonic muscle development of the grasshopper.

To analyse segmental differentiation processes in muscle development, we studied the embryogenesis of the ventral body wall muscles in thoracic and abdominal segments of the grasshopper Schistocerca gregaria at the identified cell level. We visualized differentiating muscle pioneer and muscle precursor cells by staining with a muscle-specific monoclonal antibody and with rhodamine-coupled phalloidin. Our results show that a similar pattern of serially reiterated early muscle pioneers is initially established in all segments. Subsequently, two major segmental differentiation processes occur. First, segment-specific sets of additional, later differentiating muscle pioneers are generated de novo. Second, segment-specific sets of existing early muscle precursors are eliminated through atrophy and eventual loss. These events have consequences for matching homonomy of muscles and their innervating motoneurons. Taken together, these processes in the embryo, in concert with postembryonic differentiation events, play critical roles in shaping the highly specialized muscular structures of the mature animal.

Close encounters among flying locusts produce wing-beat coupling.

Any flying animal leaves behind a wake of turbulent air. Thus, a closely tailing neighbor may be buffeted by complex aerodynamic forces. We report here that pairs of tethered locusts (Locusta migratoria) flying in tandem in a wind tunnel, couple their wing-beats to one another.Wind-receptive hairs on the rear partner's head provide the main sensory input that produces the coupling. The phase angle of coupling depends upon the distance between the individuals. By phase-coupling to a forward neighbor's wake, a locust may turn this turbulence to its own aerodynamic advantage. Moreover, within a large swarm local groups of locusts may fly in a functionally integrated manner.

Structural homology of identified motoneurones in larval and adult stages of hemi- and holometabolous insects.

The set of neurones innervating the dorsal longitudinal muscles was studied with cobalt and nickel backfills in: (1) larval and adult locusts (Schistocerca gregaria and Locusta migratoria), (2) the larval and adult beetle (Zophobas morio), and (3) various segments of these insect species. In all specimens 11 neurones were encountered, which can be subdivided into a group of 7 motoneurones that stem from the next anterior ganglion and 4 neurones located in the ganglion of the segment containing the muscles. The latter group comprises 2 contralateral and 2 medial somata, of which one is a dorsal unpaired median neurone. The results were analysed under different aspects. This neural set and the basic structure of the dendritic fields is similar in: (1) different segments (serial homology), (2) the larval stage and imago of the same species with or without a pronounced metamorphosis (ontogeny), and (3) the studied hemi- and holometabolous insects (phylogeny). Our results support the notion that the structure of these neurones is conserved irrespectively of changes in the periphery and strategy of postembryonic development.

Programmed death of peripheral pioneer neurons in the grasshopper embryo.

The Ti1 pioneer neurons arise at the distal tip of the metathoracic leg in the grasshopper embryo, and are the first neurons in the limb bud to extend axons to the central nervous system (C. M. Bate (1976) Nature (London) 260, 54-56; H. Keshishian (1980) Dev. Biol. 80, 388-397). By providing a neural pathway along which growth cones of later arising neurons migrate, these pioneer axons establish the route of one of the major nerve trunks in the leg (Keshishian, 1980; R. K. Ho and C. S. Goodman (1982) Nature (London) 297, 404-406; D. Bentley and H. Keshishian (1982) Science 218, 1082-1088). Here, we demonstrate that at the 55-59% stage of development, the two Ti1 pioneer neurons undergo programmed death. The role which these pioneers serve in establishing a nerve route appears to be their only function, and may be important for the normal development of the peripheral nervous system. The Ti1 pioneers provide an example of a previously hypothesized class (J. W. Truman (1984) Annu. Rev. Neurosci. 7, 171-188) of programmed neuron death: obsolete neurons whose function was developmental rather than behavioral.

Histological characterization of neurones innervating functionally different muscles of Locusta.

The innervation of four functionally different muscles (subalar, remotor 1, remotor 2, pleuroalar), all served by the same nerve branch, was studied in both winged segments of the locust, Locusta migratoria. Several anatomical techniques were applied: With the cobalt backfill and silver intensification technique four cell types (motoneurone, dorsal unpaired median neurone, common inhibitory neurone, and small median neurone) were demonstrated. Serial sections enabled the morphology of the motoneurones to be described in more detail and in respect to a possible functional organization of the arborizations. A differential staining technique allowed us to stain various neurones in different colours in the same preparation. With this technique the anatomy of both the "rostral" and the "caudal" subalar motoneurones could be described in parallel, thus avoiding errors in comparison due to possible individual variations from preparation to preparation. Axon counts in the peripheral nerve branch enabled us to compile a list of the total innervation for each muscle. Results from other orthopterans are integrated and whether differences in the dendritic fields might be of functional significance is discussed.

Dipteran flight motor pattern: invariabilities and changes during postlarval development.

For Calliphora the wingbeat frequency and the underlying motoneuronal activity were recorded during adult life. Wingbeat frequency increases during the ten days following last molt. The activity of motoneurons serving four selected flight muscles (nonfibrillar and fibrillar ones) also increases with age. The motoneuronal activity of young and old flies was analyzed statistically (serial and cross-correlograms, latency and phase histograms). In addition, several wing manipulations were carried out to evaluate the significance of sensory feedback on pattern generation during maturation. These ontogenetic studies suggest a centrally generated motor pattern that (1) is essentially complete with the molt to adulthood, (2) shows a progressive increase in intrinsic activity, and (3) is modulated by sensory feedback from the wing region by the same amount irrespective of age. Similarities in the postlarval development of the flight pattern of neurogenic and myogenic flyers are discussed.

Light- and electron-microscopic analysis of a complex sensory organ: the tegula of Locusta migratoria.

Structure and organization of the tegula, a cupola-shaped structure located at the anterior base of the wings of locusts, is described using various morphological methods. Based on histological and cytological criteria, two different sensory systems are distinguished: (1) a field of mechanoreceptive hairs, and (2) a chordotonal organ. The total number of sensory cells corresponds to the number of axons within the nerve supporting the tegula. The hairs are situated at the posterior region of the tegula, and each hair is innervated by only one sensory cell. The complex architecture of the chordotonal organ is analyzed and the attachment of the scolopidia to the cuticle is described. A single scolopidium makes contact with several epidermal cells. The attachment cells run in parallel and are oriented longitudinally within the tegula, being connected to each other and to the epidermal cells by desmosomes. A function in relation to wing movements during flight is suggested for the two sensory systems within the "mixed" sense organ, tegula.