Source-to-sink transport of sugar is one of the major determinants of plant growth and relies on the efficient and controlled distribution of sucrose (and some other sugars such as raffinose and polyols) across plant organs through the phloem. However, sugar transport through the phloem can be affected by many environmental factors that alter source/sink relationships. In this paper, we summarize current knowledge about the phloem transport mechanisms and review the effects of several abiotic (water and salt stress, mineral deficiency, CO2, light, temperature, air, and soil pollutants) and biotic (mutualistic and pathogenic microbes, viruses, aphids, and parasitic plants) factors. Concerning abiotic constraints, alteration of the distribution of sugar among sinks is often reported, with some sinks as roots favored in case of mineral deficiency. Many of these constraints impair the transport function of the phloem but the exact mechanisms are far from being completely known. Phloem integrity can be disrupted (e.g., by callose deposition) and under certain conditions, phloem transport is affected, earlier than photosynthesis. Photosynthesis inhibition could result from the increase in sugar concentration due to phloem transport decrease. Biotic interactions (aphids, fungi, viruses…) also affect crop plant productivity. Recent breakthroughs have identified some of the sugar transporters involved in these interactions on the host and pathogen sides. The different data are discussed in relation to the phloem transport pathways. When possible, the link with current knowledge on the pathways at the molecular level will be highlighted.

/pdf/source-to-sink-transport-of-sugar-and-regulation-by-3izapdndox.pdf

Source-to-sink transport of sugar and regulation by environmental factors

Homoptera live and feed on plants. In some circumstances, Homoptera may be tended and defended by ants; in others, they are attacked and eaten. These interactions between plants, sap-feeding Homoptera, and ants can affect each of the participants in a variety of ways. They can also modify, or be modified by, interactions between each of the participants and other organisms, such as plant pathogens, other herbivorous insects, and predators and parasites of the Homoptera. All these interactions may be influenced by abiotic factors such as soil nutrients or availability of moisture. These plant-homopteran-ant interactions are significant for several reasons. As multi-species interacting systems, they provide a link between studies of single-species life histories (156, 243) and two-species interactions (18, 103) on the one hand, and studies of overall community or ecosystem pattern and process, with the precise species undefined, on the other. They also provide a model system for studies of population genetics and evolution, as they (a) show a wide range of specificity, obJigacy, and adaptation, (b) include a variety of life histories and breeding systems, including parthenogenesis, and (c) are sensitive to genetic variation at the scale of varieties, biotypes, and individual clones. Many of the world's major plant pests are Homoptera, and many of the worst diseases of major crops are transmitted by Homoptera. Factors influenc­ ing homopteran populations and movements, and techniques to reduce them, are hence of major practical and economic significance. The manipulation· of ant assemblages to control homopteran pests has been practiced in China since

Interactions involving plants, homoptera, and ants

The role of interspecific interactions among herbivorous insects is considered to be limited, especially in specialist communities. In the current study we report on exploitative interspecific interaction between two closely related phloem—feeding species of gall—forming aphids (Homoptera; Pemphigidae; Fordinae), mediated by the supply of photoassimilates from the host plant. Geoica sp. forms a spherical gall on the leaflet midrib of Pistacia palaestina (Anacardiaceae), while Forda formicaria forms crescent—shaped galls on the leaflet margin of the same host plant. Using 14C labeling, we were able to trace the food supply (assimilated carbohydrates) from the leaves to galls of each species. We found that Geoica galls are strong sinks. These galls divert the normal phloem transport of the plant and reduce the amount of assimilates imported by F. formicaria, especially when they are located on the same leaflet. By the end of the season Geoica caused death of 84% of F. formicaria galls that were located on the same leaflet, and reduced reproductive success in the surviving galls by 20%. This is because the presence of Geoica causes early senescence (but not abscission) of the leaflet it is on (whether or not F. formicaria is present). The interaction is asymmetrical: F. formicaria did not affect reproductive output of Geoica nor did it cause visible damage to the leaflets. To our knowledge, this it the first demonstration of exploitation competition for plant assimilates between two insect—induced sinks. This exploitative competition, mediated by manipulation of plant phloem transport, stands in contrast to the absence of interference competition for galling sites between the two aphid species. Although their spatial distributions partly overlapped, the niche breadth of each species (measured from gall positions on leaves along the shoot axis) was not affected by the presence of the other. Moreover, when both species were located on the same leaf, they formed galls independently on the same or different leaflets, and there was no indication of interference competition over galling sites.

/pdf/interspecific-competition-among-phloem-feeding-insects-1puocg6tdv.pdf

Interspecific competition among phloem-feeding insects mediated by induced host-plant sinks

We investigated interactions between five species of phloem-feeding aphids (Homoptera: Aphididae) and their host plants at elevated CO2; Acyrthosiphon pisum (Harris) on Vicia faba L., Aphis nerii Boyer de Fonscolombe on Asclepias syriaca L., Aphis oenotherae Oestlund on Oenothera biennis L., Aulacorthum solani (Kaltenbach) on Nicotiana sylvestris Speg. & Comes and Myzus persicae (Sulzer) on Solanum dulcamara L. Host plants grown at elevated CO2 generally had greater biomass, leaf area and C:N ratios than those grown at ambient CO2, while plants with aphids had lower biomass and leaf area than those without aphids.



The responses of aphid populations to elevated CO2 were species-specific with one species increasing (M. persicae), one decreasing (A. pisum), and the other three being unaffected. CO2 treatment did not affect the proportion of alate individuals produced. In general, aphid abundance was not significantly related to foliar nitrogen concentration.



We performed separate analyses to test whether either aphid presence or aphid abundance modified the response of host plants to elevated CO2. In terms of aphid presence, only three of the potential 15 interactions (five aphid species x three plant traits) were significant; A. solani slightly modified the response of the plant biomass to elevated CO2 and M. persicae affected the response of leaf area and allocation. In terms of aphid abundance, only two of the potential 15 interactions were significant with A. nerii modifying the plant response to CO2 in terms of total leaf area and allocation.



We conclude that, in contrast to other insect groups such as leaf chewers, populations of most phloem-feeders may not be negatively affected by increased CO2 concentrations in the future. The reasons for this difference include the possibility that aphids may be able to compensate for changes in host plant quality by altering feeding behaviour or by synthesizing amino acids. In addition, there is little evidence that aphid herbivory, even at high levels, will substantially modify the response of plants to elevated CO2.

http://web.stanford.edu/~cbauburn/basecamp/dschool/homeproject/Hughes_CO2andAphids.pdf

Effects of elevated CO2 on five plant‐aphid interactions

The response to a single defoliation was studied on three clones of Themeda triandra collected in the short, mid, and tall grassland regions of the Serengeti National Park (Tanzania). These sites represent a gradient of decreasing grazing intensity. Growth, allocation pattern, and several morphometric traits were monitored during an 80-day period. Clipped plants of the short and medium clones fully compensated for the reduction of biomass, while plants of the tall clone showed overcompensation. During the first two weeks after clipping, clipped plants showed lower relative growth rates than unclipped ones, whereas the opposite was observed later on. Clipped plants compensated for the removal of leaf area by producing new leaves with lower specific weights and higher nitrogen content. They also produced more, smaller tillers. Although clipped plants mobilized nonstructural carbohydrates from roots and crowns, this did not account for a significant amount of growth. Relative growth rates of unclipped plants of the short clone were higher. The relative growth rate of the short clone diminished less after clipping, but also exhibited the lowest increase later. The tall clone was the most negatively affected early, but showed the highest compensation later. Compared to the other clones, the short ecotype showed many of the characteristics that defoliation induced in each individual of any clone: higher allocation to leaf area production, higher relative growth rate, higher number but smaller size of tillers, and lower leaf specific weights.

https://www.jstor.org/stable/info/4218759

Intraspecific variation in the response of Themeda triandra to defoliation: the effect of time of recovery and growth rates on compensatory growth.

Net CO2 exchange rates and dark respiration rates were determined for single attached legume leaves (leaflets) after 6 to 9 days of aphid infestation. Plant-aphid combinations used were broad bean (Vicia faba L. cv. Aquadulce) and cowpea [Vigna unguiculata (L.) Walp. cv. Caloona)] infested with cowpea aphids (Aphis craccivora Koch) and broad bean and garden pea (Pisum sativum L. cv. Victory Freezer) infested with pea aphids [Acyrthosiphon pisum (Harris)]. Leaves from all aphid-infested plants had significantly greater net CO2 exchange rates in the light than their respective controls and rates of dark respiration of leaves from infested cowpea and garden pea were also significantly greater than those of controls. Dark respiration, as a percentage of net CO2 exchange rates in the light, was greater in aphid-infested than in control plants. When the mean net daily carbon gain was calculated for the leaves of each plant-aphid combination, leaves from aphid-infested plants had the greatest gain. It is proposed that net CO2, exchange rates increased due to increased sink demand and dark respiration rates increased to meet the increased energy requirements of phloem loading and cellular maintenance associated with aphid feeding. The apparent compensatory carbon gain of infested leaves was consumed by the aphids.

Short-term effects of aphid feeding on photosynthetic CO2 exchange and dark respiration in legume leaves

The effects of various densities of cowpea aphids (Aphis craccivora Koch) and pea aphids (Acyrthosiphon pisum Harris), both Homoptera: Aphididae, on the growth of cowpea (Vigna unguiculata (L.) Walp. cv. Caloona), broad bean (Vicia faba L. cv. Aquadulce), and garden pea (Pisum sativum L. cv. Victory Freezer) seedlings were investigated. Within 10 days of infestation, aphid feeding significantly reduced plant dry weights and mean relative growth rates for the six plant–aphid combinations. In all cases except one, the mean unit leaf or net assimilation rate was also significantly reduced within 10 days. The mean leaf area ratio was the same for infested and control plants. The aphid-induced changes in host plants appear to be due to changes in photosynthesis, respiration, and translocate removal from the phloem over the 10-day period. Changes in the growth patterns of the host plant within this period are similar, but the underlying physiological effects could vary among particular plant–aphid combinations.

Aphid-induced changes in growth indices of three leguminous plants: unrestricted infestation

The short-term effects of the feeding of cowpea aphids (Aphis craccivora Koch) and pea aphids (Acyrthosiphon pisum (Harris)), both Homoptera: Aphididae, on 14C translocation and plant growth of broadbean (Vicia faba L. cv. Aquadulce), cowpea (Vigna unguiculata (L.) Walp. cv. Caloona), and garden pea (Pisum sativum L. cv. Victory Freezer) seedlings were investigated, but not all plant–aphid combinations were utilized. Within 10 days of infestation, aphid feeding reduced the flux of translocate to the roots, changed the assimilate partitioning pattern in affected shoots, and apparently induced assimilate sources to become assimilate sinks. Cowpea aphid feeding also caused more lateral branches to be formed in broadbean. Some of these effects may be related to the imbibing of translocate by aphids, while other effects may result from a series of interactions involving substances in the saliva of aphids, plant hormones, and the assimilate ratio of sources–sinks. The amount of radioactivity found per unit weig...

The effect of short-term aphid feeding on the partitioning of 14CO2 photoassimilate in three legume species

The effects of cowpea aphids (Aphis craccivora Koch) and pea aphids (Acyrthosiphon pisum (Harris)), both Homptera: Aphididae, on nitrogen (N) and phosphorus (P) uptake by and growth of cowpea (Vigna unguiculata (L.) Walp. cv. Caloona), broad bean (Vicia faba L. cv. Aquadulce), and garden pea (Pisum sativum L. cv. Victory Freezer) seedlings were investigated. After 10 days of aphid infestation, all leaf areas were significantly lower in infested plants, and plant dry weight, mean relative growth rate, and unit leaf rate were significantly lower in all plant–aphid combinations except for pea – pea aphids. The mean leaf area ratio was the same for infested and control plants indicating that infested plants did not reallocate their assimilate resources in response to aphid feeding. The accumulation of N and P as a percentage of plant dry weight did not differ between control and infested plants and was specific to the species examined. However, control plants all had greater absolute amounts of N and P after ...

Interactions between aphid infestation and plant growth and uptake of nitrogen and phosphorus by three leguminous host plants

Two-week-old cowpea seedlings (Vigna unguiculata (L.) Walp.) were infested with cowpea aphids (Aphis craccivora Koch (Homoptera: Aphididae)) for 5, 10, 15, and 20 days at which time the aphids were removed with a systemic insecticide. The seedlings were then allowed to grow for another 3 months. By day 10, plant dry weights, mean relative growth rates , and mean unit leaf rates (Ē) were all significantly reduced in the infested plants, and these relationships still held at day 20. Apparent decreases in photosynthesis and (or) increases in respiration were the primary causes of these reductions. There were no significant differences between control and experimental treatments for calculated from day 0, and total number of seeds and seed pods produced at the end of the study. Control plants had significantly more mature seed pods and seeds, while formerly infested plants had significantly more mature and unripe seeds per pod, more unripe seed pods and seeds produced, and greater values for from the time of ...

M. J. Aston

Papers

Short-term effects of aphid feeding on photosynthetic CO2 exchange and dark respiration in legume leaves

Aphid-induced changes in growth indices of three leguminous plants: unrestricted infestation

The effect of short-term aphid feeding on the partitioning of 14CO2 photoassimilate in three legume species

Interactions between aphid infestation and plant growth and uptake of nitrogen and phosphorus by three leguminous host plants

Long-term effects on cowpea plant growth of a short-term cowpea aphid infestation