Predictions & Data for this entry
| Model: hex | climate: A, BS, Cs, Cwa, Cwb, Cfa, Cfb, Dfa, Dfb, Dfc | migrate: | phylum: |
| COMPLETE = 4.5 | ecozone: TH, THp, THn, TN, TP, TPa, TA, TO, TS | food: biHl | class: |
| MRE = 0.259 | habitat: 0iTh, 0iTf, 0iTi, 0iTs, 0iTg, 0iTa, 0iTd | gender: D | order: |
| SMSE = 0.138 | embryo: T, Th | reprod: Apf | family: |
Zero-variate data
| Data | Observed | Predicted | (RE) | Unit | Description | Reference |
|---|---|---|---|---|---|---|
| Le | 0.1414 | 0.1266 | (0.1048) | cm | body length at imago stage | NevoColl2001 |
| te_MaTang | 8.63 | 6.19 | (0.2827) | d | time since birth at imago emergence | MaTang2022 |
| te_UllaGul | 6.5 | 4.679 | (0.2801) | d | time since birth at imago emergence | UllaGul2019 |
| te_DuGe1 | 5.38 | 5.029 | (0.06529) | d | time since birth at imago emergence | DuGe2004 |
| te_DuGe2 | 5.45 | 5.029 | (0.07729) | d | time since birth at imago emergence | DuGe2004 |
| te_WangZhan | 4.8 | 5.029 | (0.04766) | d | time since birth at imago emergence | WangZhan2024 |
| te_PatiPate | 8.46 | 5.292 | (0.3745) | d | time since birth at imago emergence | PatiPate2013 |
| Rj_PatiPate | 6.5 | 4.438 | (0.3172) | #/d | reproduction rate | PatiPate2013 |
| Rj_MaTang | 0.73 | 1.689 | (1.314) | #/d | reproduction rate | MaTang2022 |
| Rj_UllaGul | 1.586 | 5.684 | (2.585) | #/d | reproduction rate | UllaGul2019 |
| am_PatiPate | 23.76 | 19.29 | (0.1882) | d | average total life span | PatiPate2013 |
| t_em_MaTang | 24.56 | 15.66 | (0.3623) | d | average life span as imago | MaTang2022 |
| t_em_DuGe1 | 10.8 | 12.48 | (0.156) | d | average life span as imago | DuGe2004 |
| t_em_DuGe2 | 11.3 | 12.48 | (0.1048) | d | average life span as imago | DuGe2004 |
| t_em_WangZhan | 29.67 | 12.66 | (0.5732) | d | average life span as imago | WangZhan2024 |
| t_em_PatiPate | 16.36 | 14 | (0.1444) | d | average life span as imago | PatiPate2013 |
| Ni_MaTang | 22.77 | 23.29 | (0.02294) | # | lifetime reproductive output | MaTang2022 |
| Ni_UllaGul | 36.22 | 57.04 | (0.5748) | # | lifetime reproductive output | UllaGul2019 |
| Ni_WangZhan | 53.18 | 23.29 | (0.562) | # | lifetime reproductive output | WangZhan2024 |
| Ni_PatiPate | 46 | 56.94 | (0.2379) | # | lifetime reproductive output | PatiPate2013 |
| tR_MaTang | 15.04 | 15.49 | (0.02995) | d | reproduction period | MaTang2022 |
| tR_WangZhan | 14.04 | 12.58 | (0.1037) | d | reproduction period | WangZhan2024 |
| tR_PatiPate | 15.5 | 15.49 | (0.0006116) | d | reproduction period | PatiPate2013 |
| t1_MaTang | 2.44 | 1.529 | (0.3733) | d | duration of larval instar 1 | MaTang2022 |
| t1_UllaGul | 1.87 | 1.146 | (0.3872) | d | duration of larval instar 1 | UllaGul2019 |
| t1_DuGe1 | 1.35 | 1.242 | (0.07983) | d | duration of larval instar 1 | DuGe2004 |
| t1_DuGe2 | 1.22 | 1.242 | (0.01822) | d | duration of larval instar 1 | DuGe2004 |
| t1_WangZhan | 1.18 | 1.242 | (0.05274) | d | duration of larval instar 1 | WangZhan2024 |
| t1_PatiPate | 2.04 | 1.296 | (0.3647) | d | duration of larval instar 1 | PatiPate2013 |
| t2_MaTang | 2.11 | 1.5 | (0.2893) | d | duration of larval instar 2 | MaTang2022 |
| t2_UllaGul | 1.52 | 1.124 | (0.2606) | d | duration of larval instar 2 | UllaGul2019 |
| t2_DuGe1 | 1.35 | 1.218 | (0.09755) | d | duration of larval instar 2 | DuGe2004 |
| t2_DuGe2 | 1.3 | 1.218 | (0.06284) | d | duration of larval instar 2 | DuGe2004 |
| t2_WangZhan | 1.09 | 1.218 | (0.1177) | d | duration of larval instar 2 | WangZhan2024 |
| t2_PatiPate | 1.68 | 1.271 | (0.2434) | d | duration of larval instar 2 | PatiPate2013 |
| t3_MaTang | 2.03 | 1.538 | (0.2424) | d | duration of larval instar 3 | MaTang2022 |
| t3_UllaGul | 1.2 | 1.153 | (0.03948) | d | duration of larval instar 3 | UllaGul2019 |
| t3_DuGe1 | 1.31 | 1.249 | (0.04625) | d | duration of larval instar 3 | DuGe2004 |
| t3_DuGe2 | 1.48 | 1.249 | (0.1558) | d | duration of larval instar 3 | DuGe2004 |
| t3_WangZhan | 1.09 | 1.249 | (0.1463) | d | duration of larval instar 3 | WangZhan2024 |
| t3_PatiPate | 2 | 1.304 | (0.3482) | d | duration of larval instar 3 | PatiPate2013 |
| t4_MaTang | 2.15 | 1.623 | (0.245) | d | duration of larval instar 4 | MaTang2022 |
| t4_UllaGul | 1.91 | 1.257 | (0.3419) | d | duration of larval instar 4 | UllaGul2019 |
| t4_DuGe1 | 1.37 | 1.319 | (0.03737) | d | duration of larval instar 4 | DuGe2004 |
| t4_DuGe2 | 1.45 | 1.319 | (0.09048) | d | duration of larval instar 4 | DuGe2004 |
| t4_WangZhan | 1.44 | 1.319 | (0.08416) | d | duration of larval instar 4 | WangZhan2024 |
| t4_PatiPate | 1.64 | 1.421 | (0.1333) | d | duration of larval instar 4 | PatiPate2013 |
| t0R_MaTang | 0.92 | 0.6745 | (0.2668) | d | adult prereproduction period | MaTang2022 |
| t0R_PatiPate | 0.68 | 0.6543 | (0.03779) | d | adult prereproduction period | PatiPate2013 |
| Le_hd | 0.0301 | 0.03146 | (0.04531) | cm | head width at imago emergence | NevoColl2001 |
| Lb_PatiPate | 0.051 | 0.07103 | (0.3927) | cm | body length at birth | PatiPate2013 |
| L2_PatiPate | 0.079 | 0.08193 | (0.0371) | cm | body length of instar 2 | PatiPate2013 |
| L3_PatiPate | 0.114 | 0.09425 | (0.1733) | cm | body length of instar 3 | PatiPate2013 |
| L4_PatiPate | 0.139 | 0.1088 | (0.2172) | cm | body length of instar 4 | PatiPate2013 |
| Le_PatiPate | 0.168 | 0.1273 | (0.2425) | cm | body length at imago stage | PatiPate2013 |
| Lb_RaniBora | 0.051 | 0.071 | (0.3922) | cm | body length at birth | RaniBora2015 |
| L2_RaniBora | 0.07 | 0.0819 | (0.17) | cm | body length of instar 2 | RaniBora2015 |
| L3_RaniBora | 0.085 | 0.09422 | (0.1084) | cm | body length of instar 3 | RaniBora2015 |
| L4_RaniBora | 0.116 | 0.1088 | (0.06232) | cm | body length of instar 4 | RaniBora2015 |
| Le_RaniBora | 0.129 | 0.1266 | (0.01877) | cm | body length at imago stage | RaniBora2015 |
| Le_alate | 0.11 | 0.1108 | (0.007476) | cm | body length at imago stage | RaniBora2015 |
| Bb_PatiPate | 0.038 | 0.03914 | (0.03009) | cm | breadth at birth | PatiPate2013 |
| B2_PatiPate | 0.047 | 0.04515 | (0.03931) | cm | breadth of instar 2 | PatiPate2013 |
| B3_PatiPate | 0.059 | 0.05194 | (0.1196) | cm | breadth of instar 3 | PatiPate2013 |
| B4_PatiPate | 0.071 | 0.05996 | (0.1554) | cm | breadth of instar 4 | PatiPate2013 |
| Be_PatiPate | 0.089 | 0.07013 | (0.212) | cm | breadth at imago emergence | PatiPate2013 |
| Bb_RaniBora | 0.025 | 0.03913 | (0.5652) | cm | breadth at birth | RaniBora2015 |
| B2_RaniBora | 0.038 | 0.04514 | (0.1878) | cm | breadth of instar 2 | RaniBora2015 |
| B3_RaniBora | 0.053 | 0.05192 | (0.02031) | cm | breadth of instar 3 | RaniBora2015 |
| B4_RaniBora | 0.067 | 0.05994 | (0.1053) | cm | breadth of instar 4 | RaniBora2015 |
| Be_RaniBora | 0.07 | 0.06976 | (0.003454) | cm | breadth at imago emergence | RaniBora2015 |
| Be_alate | 0.06 | 0.06107 | (0.01791) | cm | breadth at imago emergence of alate | RaniBora2015 |
Uni- and bivariate data
| Data | Figure | Independent variable | Dependent variable | (RE) | Reference |
|---|---|---|---|---|---|
| T_Re_ZamaTale |   | temperature | reproduction rate | (0.4075) | ZamaTale2006 |
| T_Re_XiaWerf | | temperature | reproduction rate | (0.3639) | XiaWerf1999 |
| T_tj_KersSata |   | temperature | time since birth at imago emergence | (0.08495) | KersSata1999 |
| T_tj_ZamaTale | | temperature | time since birth at imago emergence | (0.1561) | ZamaTale2006 |
| T_tj_XiaWerf | | temperature | time since birth at imago emergence | (0.1718) | XiaWerf1999 |
| T_t_em_XiaWerf |   | temperature | average lifespan as imago | (0.3559) | XiaWerf1999 |
| T_t_em_ZamaTale | | temperature | average lifespan as imago | (0.7097) | ZamaTale2006 |
| T_t1_XiaWerf |   | temperature | duration of 1st larval instar | (0.1682) | XiaWerf1999 |
| T_t1_ZamaTale | | temperature | duration of 1st larval instar | (0.07068) | ZamaTale2006 |
| T_t2_XiaWerf |   | temperature | duration of 2nd larval instar | (0.1815) | XiaWerf1999 |
| T_t2_ZamaTale | | temperature | duration of 2nd larval instar | (0.2037) | ZamaTale2006 |
| T_t3_XiaWerf |   | temperature | duration of 3rd larval instar | (0.1624) | XiaWerf1999 |
| T_t3_ZamaTale | | temperature | duration of 3rd larval instar | (0.2444) | ZamaTale2006 |
| T_t4_XiaWerf |   | temperature | duration of 4th larval instar | (0.1842) | XiaWerf1999 |
| T_t4_ZamaTale | | temperature | duration of 4th larval instar | (0.1665) | ZamaTale2006 |
| T_am_XiaWerf |   | temperature | average total lifespan | (0.1425) | XiaWerf1999 |
| T_am_KersSata | | temperature | average total lifespan | (0.2304) | KersSata1999 |
| tN_MaTang |   | time since birth | cumulative reproduction | (0.1751) | MaTang2022 |
| tN_UllaGul | | time since birth | cumulative reproduction | (0.3655) | UllaGul2019 |
| tN_ZamaTale15 |   | time since birth | cumulative reproduction | (0.2958) | ZamaTale2006 |
| tN_ZamaTale20 | | time since birth | cumulative reproduction | (0.5248) | ZamaTale2006 |
| tN_ZamaTale25 | | time since birth | cumulative reproduction | (0.3636) | ZamaTale2006 |
| tN_ZamaTale30 | | time since birth | cumulative reproduction | (0.6034) | ZamaTale2006 |
| T_t0R |  | temperature | duration of prereproductive period | (0.3723) | XiaWerf1999 |
| tS |  | time since birth | survival rate | (0.2725) | MaTang2022 |
| tWw |  | time since birth | wet weight | (0.2027) | Gao2019 |
Pseudo-data at Tref = 20°C
| Data | Generalised animal | Aphis gossypii | Unit | Description |
|---|---|---|---|---|
| v | 0.02 | 0.006708 | cm/d | energy conductance |
| p_M | 18 | 33.11 | J/d.cm^3 | vol-spec som maint |
| k_J | 0.002 | 0.002 | 1/d | maturity maint rate coefficient |
| k | 0.3 | 0.2658 | - | maintenance ratio |
| kap | 0.8 | 0.8 | - | allocation fraction to soma |
| kap_G | 0.8 | 0.8083 | - | growth efficiency |
| kap_R | 0.95 | 0.95 | - | reproduction efficiency |
| s_s | 0.01 | 5.917e-09 | - | supply stress |
Discussion
- This entry was developed based on the entry for Myzus persicae
- For now, this entry focuses only on apterous viviparous females of the summer cycle, i.e., the most common form used in laboratory studies. Zoom factor for alates is based on a single observation.
- Investment into reproduction starts at birth, thus the hex model was used instead of abp (approach adopted from Myzus persicae)
- The pupa phase of the model type hex is skipped, so that the event j indicates the final moult to the imago stage instead; consequently, parameters kap_V and E_He remain unused
- Imagos are assumed to behave like in model type abp: growth stop, no kappa rule, continued feeding and reproduction
- The release rate of offspring is assumed to match the repro investment of the imago 1:1, so d_ER is net zero, and ERj remains as a buffer (see next discussion point)
- All repro investment before the imago stage serves as a time-buffer to have fully developed embryos ready by the end of the preoviposition period
- The preoviposition period is covered by a new parameter t0R ("preoviposition period at reference temperature"), which should be corrected for temperature but which for now is independent of f; Reproduction period is defined as the time from first to last oviposition
- An upper temperature boundary was used to capture reduced development and reproduction around 30 C; total lifespan generally does not follow this pattern
- Parameter kap was fixed at 0.8 to avoid convergence to 1; data on repro buffer at final moult would be needed to avoid this
- No maternal effects were assumed when calculating egg costs at different f-levels; otherwise, f-values for different food sources might not have the intended effect on reproduction rates
- Discrepancy in T-Rj curves across studies is comparable to coefficient variation in XiaWerf data - difference might not be meaningful
- "Breadth" is interpretable as total body breadth; no further details on breadth measurements available
- Available reproduction data from KersSata was excluded from the fitting because the reported Rj values could not be replicated; also, the temperature-dependency of reproduction could not be integrated with that from other studies
- It is possible to achieve at parameter values for which adult pre-reproductive period and reproductive period exceeds the lifespan as imago, which is not plausible; when updating the fit, it is advisable to check that the difference (t_0R + tR) - t_am is acceptable
- A better fit to T-am data could be achieved by reducing the initial value for h_a, but this could lead to implausible parameter values (see previous point)
- Temperature curves do not match total lifespan at high temperatures, possibly indicating some additional heat stress that is not accounted for through basic temperature correction; this is currently ignored
- Maximum length was always taken to be Le, not Lm (which is never reached). Correspondingly, the aging module was adapted for the hex model so that s_G is linked to (L/L_e)^3 instead of (L/L_m)^3
- Shape of the survival curve does not match sigmoid curve predicted by standard aging submodule, suggesting some additional cause of mortality or life stage-specific mortality; the standard aging module was used
- With the current model formulation, the temperature response of Ni will not be matched correctly; Achieving this will require more development work, especially regarding the mechanisms behind the reproductive plateau; the default hex model does not cover this since N lugens imagos do continue to feed
- Reproduction rates peak in the early adult phase and go to 0 with increasing age; this is currently ignored, and only the initial (linear) reproduction phase is fitted
- For some studies, the model is simultaneously fitted to zerovariate (Rj, tj, Ni) and univariate data (tN), effectively increasing the weight for these observations; after testing different options, this led to the best overall description of the data
- Weights for tWw and Rj_ZamaTale were increased; we found this to lead to an overall more balanced fit
- Observed tS does not appear to follow the sigmoid trajectory implied by the standard aging module, suggesting different, possibly life stage-specific, sources of mortality
- Particularly high reproduction rate observed in PatiPate2013 (food: Plantago ovata); the reasons are not clear, other data types do not show congruent differences
Facts
- Mostly asexual reproduction, although sexual reproduction also ocurrs (Ref: Eber1997)
- Nymphs reach imago in 4 molts with no pupa stage (Ref: Eber1997)
- Feeding continues in imago stage; e.g. Liu et al. (2021) studied feeding behaviour in adults (Ref: LiuJin2021)
- Aphis gossypii is cosmopolitan, with a preference for warmer regions (Ref: forest)
- In Aphids, investment in reproduction likely occurs very early in development (Ref: Akim2006)
Acknowledgment
- The creation of this entry was financially supported by Bayer AG
Bibliography