Hello,

This review also contains info relevant to this discussion: 
https://doi.org/10.1016/j.sbi.2019.04.006

Figure 1 shows more examples of electron versus x-ray atomic scattering factors 
for various elements and their charged states.

And this paragraphs is very relevant to Matthew's questions:


"X-ray crystallographic electron density maps accurately represent the 
positions of atomic centers because peaks in most maps are centered on the 
nuclei. This approximation breaks down at very high resolution (<1 Å) where 
electron orbital structure becomes resolved and electron density peaks are 
distributed around nuclei [11]. In contrast, peaks in Coulomb potential maps [= 
cryoEM maps] are not necessarily centered on atomic nuclei, particularly when 
atoms carry a net charge. Unlike X-ray scattering, electron scattering has a 
strong charge dependence (e.g. compare O and O− in Figure 1a). This phenomenon 
can result in deviations of Coulomb potential map features from atomic centers 
that can confound accurate model building using current methods."


So, it seems like difficult agreement between a cryoEM map and ideal 
stereochemistry is to be expected.


The review goes on to cite a series of papers discussing this point in more 
details:


https://doi.org/10.1002/pro.3060

https://doi.org/10.1002/pro.3169

https://doi.org/10.1002/pro.3198 : this paper explains how to convert a Coulomb 
potential map (= an unfiltered/unsharpened cryoEM map) into a charge density 
map, which is presumably easier to interpret and in better agreement with ideal 
geometry? I have not yet tried this, and in fact have not yet read this series 
of papers closely. But this seems very interesting, I will try this with the 
atomic resolution apoF cryoEM maps from 2020 if/when I find the time.

https://doi.org/10.1002/pro.3093


I hope this helps,


Guillaume

________________________________
From: CCP4 bulletin board <[email protected]> on behalf of zbyszek 
<[email protected]>
Sent: Tuesday, July 29, 2025 1:56:18 AM
To: [email protected]
Subject: Re: [ccp4bb] Restraining metal sites in medium resolution cryo-EM


I believe much more work is needed to understand charge effects in cryoEM. An 
interesting paper (https://www.nature.com/articles/s42004-023-00900-x) shows in 
Fig. 3 that these effects for oxygen extend to medium resolution. However, a 
complete understanding also requires consideration of radiation effects, as 
radiation exposure rapidly redistributes charges in addition to causing damage 
and decay.

The charge on a given atom may already be affected by exposures in the kGy 
range, meaning that even the first frame of a cryoEM movie is impacted. The 
first frame typically corresponds to a dose of ~3–4 MGy (~1 e⁻/Ų). Total doses 
in cryoEM are high (on the order of hundreds of MGy) so radiation-induced 
damage to sensitive groups, including those that coordinate metal cations, is 
significant. Because a cryoEM structure, even after up-weighting the early 
frames, represents a partially damaged state, the coordination sphere, if left 
unrestrained, is difficult to interpret.

Zbyszek


On 2025-07-28 18:31, James Holton wrote:

Yes, but formal charge effects are not implemented.  Last time I checked?

-James Holton
MAD scientist

On 7/25/2025 11:01 AM, Dominika Borek wrote:

I don't think there's anything particularly difficult about scattering factors 
for cryoEM, but the patterns of dependences on atomic number differs from that 
in X-ray crystallography. For instance, in X-ray maps, oxygen atoms, common 
interactors of metal cations via carboxylic acids and/or water molecules, 
appear "stronger" due to this difference. Thus, at the same nominal resolution 
of ~2–3 A, the coordination sphere in cryoEM maps looks less well-defined than 
in X-ray crystallography maps (even though information content is the same). I 
find the default restraints insufficient, so I apply restraints manually in 
both Refmac/Servalcat and Phenix, e.g., I define Mg2+ octahedral geometry with 
proper distances in a mini script and include it in the refinement run. In 
other words, I vote for tighter geometry.



D.



Dominika Borek, Ph.D. *** UT Southwestern Medical Center

5323 Harry Hines Blvd. *** Dallas, TX 75390-8816

214-645-9577 (phone) *** 214-645-6353 (fax)







From: CCP4 bulletin board <[email protected]><mailto:[email protected]> 
On Behalf Of Frank von Delft
Sent: Friday, July 25, 2025 10:10 AM
To: [email protected]<mailto:[email protected]>
Subject: Re: [ccp4bb] Restraining metal sites in medium resolution cryo-EM



Isn't there something about scattering factors being ... difficult... for 
CryoEM reconstructions?

Which I imagine would cause issues in refinement.

Frank


On 25/07/2025 12:35, Matthew Snee wrote:

Hi Everyone



Id let to get the communities opinions on metal sites in Cryo-EM models.



In X-ray structures the geometry of metal coordination (when restrained 
correctly) looks pretty good, even in low resolution structures, and at very 
high resolution you may wish to avoid applying restraints so that the precise 
coordination distances/angles can tell you something about the physical 
chemistry of the system (I.E reduction state of catalytic metals in chemical 
biology).   I have found that EM structures between 2-3Å which are good enough 
to see individual features (un ambiguous sidechain rotamers and coordinated 
waters in the metal complex) refinement outputs don't look nearly as nice.



My question is this,:



Is this just "the way it is" because the optical resolution and true atomic 
resolvability is different between X-ray and EM.

The constraints of the crystal might help to collapse the conformational 
landscape somewhat, and solvent flattening might repress (unidentifable) minor 
states that would distort the real-space fit, thus improving the convergence 
between ideal geometry and fit to the density.

EM structures that have very high resolution FSC cutoffs can still be distorted 
by minor anisotropic flexibility in one or more particular direction,  there 
are ways to isotropise maps, but I dont really like refining against these.



The other option is that the restraints need to be up-weighted so that they 
have more parity with the protein bond restraints and other targets that one 
might use.



Basically, are these structures where the coordination geometry is almost 
certainly not completely correct, likely to be better, more useful, or closer 
to the "true structure" or is it better to enforce the geometry more strictly 
than you would for an equivalent X-ray structure in order to achieve the most 
likely sensible cluster.



I dont think either approach is "wrong" because you could make an argument for 
building what you see, only building the links that are directly suggested and 
restraining them as best you can,  I'm not a metal cluster expert by any means, 
but I do know that the coordination number, and bond length/angles are quite 
well known.



I know this is potentially more of a CCPEM/Phenix query, but X-ray people are 
generally more involved in the physical chemistry side (I know not always 🙂!),



Best wishes





Matthew Snee, PhD

Post-doctoral Research Associate

The Baldock Lab | Michael Smith Building C3.214 | Wellcome Trust Centre for 
Cell-Matrix Research |Division of Cell Matrix Biology & Regenerative Medicine| 
School of Biological Sciences| Faculty of Biology, Medicine and Health, 
University of Manchester, Oxford Road, Manchester M13 9PT

Lab Tel: +44 (0)161 306 2869



[cid:[email protected]][cell-matrix-research]





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